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TORREYA

A Bi-MonTHLY JOURNAL oF BotranicaL Notes Anp News

John Torrey, 1796-1873

EDITED FOR
THE TORREY BOTANICAL CLUB
BY
WEEE |) BO NTSAMs EE
AND

HAROLD EO LUM
VOLUME 42

New York
1942

sng Pia.

NEW YORK
ANICAI

Volume 42 January-February, 1942 Number I

TORREYA

A Bi-MontTHLYy JoURNAL oF Botanica, Notes AND NEws

GARDEN

EDITED FOR
THE TORREY BOTANICAL CLUB

WILLIAM J. BONISTEEL

John Torrey, 1796-1873

CONTENTS

Thomas Horsfield—American Naturalist and Explorer....... James B. McNair 1
Notes on the Flora of Arizona...............-202e-eeeacecees LyMAn BENSON
Dhe7Names of Gormusijs 325 c cic ots ais Wales se shelelore cue sls cieiol eter ckoush H. W. Rickert I1
Phyllanthus nummulariaefolius Poir. in the United States........ Lron Croizat 14
Book Reviews

Ane Jee \WWard tls coving dos moa scacdom acc Aueues ca ade oa Greorce T. Hastines 18

ThesAdvance ofthe Bungie. 95sec se sacle erie ses eaten = etal tek B. O. Donce 19

Hunger Signs in Crops.............2-20 00sec e eee eee Wo. J. BonisTEEL 21
Field ‘Trips of the Clubs.) .2210.. 2 foe. onc ae te ce gs rierlege ee minim = cilia 22
Proceedings of the Club.............. BN cata le em LON REDE Seah ets a cat al ace a teres 25
ING ws INO CG oar ee ie eMC IN Sree corres aa et Val viataDa\talol efava(eskel » 9/a\)) aletereieis etn fens ee19 32

PUBLISHED FOR THE CLUB

By THE FREE PRESS PRINTING COMPANY
187 CoLLEGE STREET, BURLINGTON, VERMONT

Entered as second class matter at the post office at Burlington, Vermont,
October 14, 1939, under the Act of March 3, 1879

TORREYA

TorreYA, the bi-monthly publication of the Torrey Botanical Club, was established
in 1901. TorrREYA was established as a means of publishing shorter papers and inter-
esting notes on the local flora range of the club. The proceedings of the club, book
reviews, field trips and news notes are published from time to time. The pages of
TORREYA are open to members of the club and others who may have short articles
for publication.

ToRREYA is furnished to subscribers in the United States and Canada for one
dollar per year (January-December); single copies thirty cents. To subscribers
elsewhere, twenty-five cents extra, or the equivalent thereof. Postal or express
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dressed to the treasurer, Dr. W. Gordon Whaley, Barnard College, Columbia Uni-
versity, New York, N. Y.

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subscription to TORREYA.

INSTRUCTIONS TO CONTRIBUTORS

The manuscript should be prepared so that it conforms to the best practice as
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TorrREYA is edited for the Torrey Botanical Club by

WM. J. BONISTEEL
W. H. CAMP DOROTHY J. LONGACRE

MEMBERSHIP IN THE TORREY BOTANICAL CLUB

All persons interested in botany are invited to join the club. There are four
classes of membership: Sustaining, at $15.00 a year; Life, at $100; Annual, at $5.00
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are: (a) To attend all meetings of the club and to take part in the business, and
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Manuscripts for publication, books and papers for review, reports of field
trips and miscellaneous news items should be addressed to:

DR. WM. J. BONISTEEL
BIOLOGICAL JLLABORATORIES, FORDHAM UNIVERSITY
New York, N. Y.

TORREYA

Vol. 42 January-February, 1942 No. 1

Thomas Horsfield—American Naturalist and Explorer

James B. McNair

The eighty-six years of Thomas Horsfield’s life may be divided
into three periods—the American period of twenty-six years from
1773 to 1799, the Javan period of twenty years from 1799 to 1819,
and the British period of thirty-nine years from 1820 to 1859. But
before taking up Dr. Horsfield’s career in detail it might be well to
devote some time to a study of his ancestry.

Thomas Horsfield’s grandfather was Timothy Horsfield’ who
was born in Liverpool, England in 1708 and was educated in the
parish school. In 1725 he emigrated to New York and joined his
brother Isaac, with whom he learned the trade of butcher. In 1735
they leased two stands in the Old Slip Market where their business
became large and profitable.

Although a member of the Church of England, he became inter-
ested in the Moravian Church in 1739. In 1748 he applied to the
authorities at Bethlehem, Pennsylvania for permission to reside
there, but because he was one of the executors of the estate of
Thomas Noble, a prominent merchant of New York, and a member
of the newly organized Moravian congregation, as well as being
entrusted with the building of the Irene, he was requested to post-
pone his removal. He, however, took his children to Bethlehem to be
educated in the schools. The year following he moved there him-
self where, except for a short sojourn in Nazareth, Pennsylvania,
he resided until his death.

On the founding of Northampton County in 1752, Timothy
Horsfield was appointed a justice of the peace by Governor Hamil-
ton. In 1763 he was commissioned colonel of the forces in the county
for the defense of its frontiers against Indian raids. This appoint-

1 Timothy Horsfield, perhaps the great grandfather of T. Horsfield, ap-
pears in the parish register of St. Nicholas Church, Liverpool in 1694 and
1704.

Torreya for January-February (Vol. 42: 1-32) was issued February 27, 1942.
1

ment excited jealousy, so he soon resigned and lost his justiceship
in consequence. Squire Horsfield lived in what was known as the
Oerter house, which stood on Market Street opposite the graveyard.

In 1731 Timothy Horsfield was married to Mary, daughter of
John Doughty, a prominent butcher of Long Island, and a lineal
descendant of the Reverend Francis Doughty, who, in 1632,
preached the first Presbyterian sermon. Both Timothy and Mary
Horsfield died in 1773 on Long Island.

Thomas Horsfield’s best known uncle was Joseph Horsfield
who was chosen a delegate to the Pennsylvania convention to ratify
the Federal Constitution in 1787 and one of the signers of the rati-
fication. In 1792 he was appointed by President Washington to be
the first postmaster of Bethlehem.

Thomas Horsfield’s father was Timothy Horsfield, Jr., who
married Juliana Parsons at Philadelphia in 1738. She was the
daughter of William Parsons, surveyor general and founder of
Easton, Pennsylvania. Timothy Horsfield died April 11, 1789 and
his wife died January 17, 1808.

Thomas Horsfield was born at Bethlehem, Pennsylvania, May
12, 1773. He received his early education in the Moravian schools
at Bethlehem and Nazareth. Very early in life his tastes led him
to the study of botany, and a similar inclination to the pursuit of
all branches of biological science may have caused him to select
medicine as a profession. He pursued a course in pharmacy with
Dr. Otto of Bethlehem and devoted special attention also to botany.
This Dr. Otto was probably John Frederick Otto, M.D., of Halle
who arrived from Europe in 1750. He was widely known as physi-
cian and surgeon and died at.Nazareth in 1779.

Thomas Horsfield graduated in medicine at the University of
Pennsylvania in 1798 in the twenty-fifth year of his age and served
as “medical apprentice” in the Pennsylvania Hospital from 1794 to
1799. While at the University he was a pupil of Dr. Benjamin
Smith Barton. “His graduation thesis is remarkable for its pains-
taking clinical description of the toxic symptoms of the poisoning _
produced by sumac and poison ivy, and for the record of well-
conceived experiments carried out upon himself and upon animals
concerning the pharmacological action of this interesting poison.
It ranks as a pioneer contribution in the history of experimental
pharmacology in America.”

The year after his graduation, in October 1799, he accepted
service as surgeon on the China, a merchant vessel about to sail for
Java. In the course of the voyage he visited Batavia, in the island
of Java. He was impressed with the beauty of the scenery, the rich-
ness of the vegetation, and certain drugs in common use by the
natives which were extracted from local plants. He decided to in-
vestigate these substances, so upon his return home he secured
such books, scientific instruments and materials as he could get
together in Philadelphia and undertook a second voyage to Batavia
in 1801. There he secured, upon application, an appointment as
surgeon in the Dutch Colonial Army, and this gave him an oppor-
tunity to visit and study the flora, fauna and geology of the various
parts of the island. This was the beginning of eighteen years of
study which linked his name inseparably with the natural history
and especially the botany of Java.

In the prefaces of his various works he tells the story of his
collections and travels. It appears that between 1802 and 1811 his
facilities were poor and many of his most valued specimens decay-
ed owing to inadequate preservation. or several years his re-
searches were confined to the vicinity of Batavia, but beginning
with 1804 he visited nearly all parts of Java and made brief trips to
several of the neighboring islands.

In 1811 Java became a British possession, administered by the
East India Company. The temporary commissioner authorized
Horsfield to continue his investigations along the same lines as
hitherto, and before the end of the year a new governor, Sir Thomas ©
Stamford Raffles (after whom the genus Rafflesia and family Raffle-
siaceae were named) confirmed his appointment in the service of
the East India Company. This connection enabled him to pursue
his studies on a more elaborate scale. Dr. Horsfield thoroughly ex-
plored every part of the island in quest of its natural products. From
Java he visited Banca and gave the fullest and best account which
exists of the mineralogy, geology, botany and zoology of that island.
After the restoration of Java to the Dutch in 1816, Dr. Horsfield
made a long sojourn in Sumatra and there continued his favorite
studies.

He secured the warm friendship of Sir Stamford Raffles, who,
it is believed, acquired from Dr. Horsfield that love of natural his-
tory: by which he was distinguished, and which rendered him so

zealous in its promotion. Dr. Horsfield followed that eminent man
to England in 1818 and soon after was made Keeper of the Museum
of the East India Company, which charge he held until his death
on July 24, 1859 in the eighty-seventh year of his age.

In regard to Dr. Horsfield’s work in Java, Sir Stamford Raf-
fles says in his History of Java that “For all that relates to the
natural history of Java, I am indebted to the communications of
Dr. Thomas Horsfield. Though sufficient for my purpose, it forms
but a scanty portion of the result of his long and diligent researches
on the subject.”

It is not strange that one who graduated in medicine and whose
graduation thesis should be a study of the action of a poisonous
plant should be interested in other plants of pharmacological action.
And so we find that upwards of sixty of the medicinal plants of
Java were described for the first time by Dr. Horsfield in the
Batavian Transactions. One of these studies which gained especial
notice was his work on the Upas tree in which he refuted the false-
hoods and fabulous traditions which had been published concerning
this subject.

Sir Stamford Raffles also states that “Upwards of a thousand
(Javanese) plants are already contained in the herbaria of Dr.
Horsfield, of which a large proportion are new to the naturalist.”
This extensive collection was sent to England and later (1858)
presented by the East India Company to the Linnean Society of
London. A selection only of his botanical collections was published
as a monograph “Plantae Javanicae Rariores.” This is a beautifully
illustrated work, prepared with the assistance of the botanists Robert
Brown and J. J. Bennett. In it 2,196 species are described, all of
which Horsfield had collected himself.

Dr. Horsfield although eminent as a botanist and equally versed
in mineralogical knowledge, was perhaps most eminent as a
zoologist. The most important of his zoological publications and
the earliest of his independent works after his coming to England,
was his “Zoological Researches in Java and the Neighbouring _
Islands,” published in 1821 and the following years. His other
zoological writings are chiefly the valuable illustrated catalogues of
mammals, birds and lepidoptera of the several zoological depart-
ments of the East India Company’s museum, and numerous papers
on zoological subjects contributed to the “Linnean Transactions.”

the “Zoological Journal” and the “Proceedings of the Zoological
Society.” His latest publication was the “Catalogue of the Lepi-
dopterous Insects in the East India Museum.” It was compiled
by Mr. Moore, his assistant, from Dr. Horsfield’s materials and
manuscripts, and under his direction. Dr. Horsfield had some years
before commenced a catalogue of these insects, of which only two
parts were published (1828-29). This publication, though incom-
plete, deserves notice, as it contains an elaborate introduction, with
a general arrangement of the Lepidoptera founded on their meta-
morphosis. The importance of the transformations of insects in
reference to their classification had indeed become early impressed
on Dr. Horsfield’s mind. He accordingty spent three seasons dur-
ing his stay in Java in collecting the larvae of numerous species of
Lepidoptera, watching their development, and making careful
descriptions and drawings of their successive changes up to the
perfect state.

Dr. Horsfield always took the deepest interest in the progress
of natural history, and especially in the systematic arrangement
of animals, in which he adopted the views of Mr. McLeay. His
classification of the diurnal lepidoptera and of birds exhibits great
powers of philosophical analysis.

His numerous scattered papers, if put together, would constitute
several large and valuable volumes, and many of them, more espe-
cially those on geology and natural history of the Eastern Archi-
pelago, well deserve to be collected in a separate form.

Dr. Horsfield was a man of retiring habits, but of amiable char-
acter and unblemished integrity. He was one of the few Americans
who became a Fellow of the Royal Society of London (in 1828).
He was a member of many other societies including the Batavian
Society, the Zoological Society of London and the Geological Society
of London. He was elected a Fellow of the Linnean Society in 1820
and later became one of its vice-presidents.

Three genera of plants have been named Horsfieldia at different
times. Horsfieldia of Willdenow (1805) is the oldest and com-
prises plants of the Myristicaceae. It is in current use and included
more than fifty species of nutmegs. The genus Horsfieldia of Blume
(1830) was composed of a species of the Araliaceae. Chifflot (1909)
designated the genus Horsfieldia for some of the Gesneriaceae.
Because Horsfieldia was first used by Willdenow in a generic sense

the genus Horsfieldia of Blume was changed to Harmsiopanax
Warb: and that of Chifflot to Monophyllaea Reichb. Many species
of plants and insects also bear Horsfield’s name.

References

Anonymous. 1859. Obituary. The Times. London. July 29, 1859.

. 1859. Obituary. Proceedings of the Royal Society of London. 10: xix—

XX1.

. 1861. Obituary. Proceedings of the Linnean Society of London. Anni-

versary meeting May 24, 1860. Jour. of the Proc. of the Linn. Soc. Zoology.

5: XXV-XXVI.

. 1858. Proceedings of the Linnean Society of London. Meeting of Nov.
4, 1858. Jour. of the Proc. of the Linn. Soc. Zoology. 4: i.
Candolle, A. P. de 1830. Prodomus systematis naturalis regni vegetabilis, sive
Enumeratio contracta ordinum, generum specierumque plantarum. 4: 87.
Carson, J. 1869. A history of the medical department of the University of
Pennsylvania. pp. 131-132.

Chifflot, J. B. J. 1909. Sur quelques variations du Monophyllaea Horsfieldii R.
Br. Compt. rend. Acad. Sci. Paris. 148: 939-941.

Egle, W. H. 1887. The Federal Constitution of 1787. The Pennsylvania Mag.
of Hist. and Biogr. 11: 213-222.

Gray, Asa. 1859, 1860. Amer. Jour. of Science. Second series (1859) 28: 444;
(1860) 29: 441.

Jordan, J. W. 1896. Timothy Horsfield. Notes and Queries. Edited by W. H.
Egle. Harrisburg, Pa. 3: 166-168. Reprinted edition.

. 1909. William Parsons, Surveyor General and Founder of Easton, Pa.
The Pennsylvania Mag. of Hist. and Biogr. 33: 345-346.

Levering, J. M. 1903. A History of Bethlehem, Pennsylvania. p. 171.

McNair, J. B. 1923. Rhus dermatitis, its pathology and chemotherapy. Univer-
sity of Chicago Press. pp. 83, 101, 116, 119, 126, 127, 139, 141, 142, 145, 147,
193.

Oliver, D. 1860. Notes on the British Herbarium of the Linnean Society. Jour.
of the Proc. of the Linn. Soc. Botany. 4: 194-198.

Raffles, Sir T. S. 1817. The History of Java. London. Vol. 1.

Willdenow, C. L. 1805. Species plantarum. 4: 872.

Publications

Horsfield, Thomas. 1798. An experimental dissertation on the Rhus vernix,
Rhus radicans and Rhus glabrum; commonly known in Pennsylvania by the -
names of poison-ash, poison-vine and common sumach. By Thomas Hors-
field of Bethlehem, Pennsylvania. Philadelphia. Printed by Charles Cist,
No. 104 North Second Street. Published also in Charles Caldwell’s “Medi-
cal Theses,” Philadelphia 1805. pp. 113-163.

. 1805. An account of a voyage to Batavia in the year 1800. The Phila-

delphia Medical Museum. Edited by J. Redman Coxe. Vol. 1.

——. 1814. Scheikundige ontleding van een vulkaansch zand en een yzer-erts.
(Chemical analysis of some volcanic sand and an iron ore.) Batav.
Genootsch. Verhand. 7.

. 1814. Over de Rivier van Solo. (Beyond the River Solo.) Batay.

Genootsch. Verhandl. 7.

. 1814. Reis naar de Ooster-streken van Java. (Journey to the East-

stretch of Java.) Batav. Genootsch. Verhandl. 7.

. 1814. Beschrijving van den Gatip-Boom. (Description of the Gatip-

Tree.) Batav. Genootsch. Verhandl. 7.

. 1814. Beknopte beschrijving van het Crinum asiaticum. (Brief descrip-

tion of Crinum asiaticum.) Batav. Genootsch. Verhandl. 7.

. 1814. Scheikundige ontleding der vruchten van den Rarak-Boom.

(Sapindus saponaria, Linn.) (Chemical analysis of the fruit of the Rarak-

Tree.) Batav. Genootsch. Verhandl. 7.

. 1814. On the Oopas or Poison-tree of Java. Batav. Genootsch. Ver-
handl. 7; Thomson Ann. Phil. 9: 202-214, 265-274 (1817) ; Jour. de Phys.
84: 259-266 (1817); Amer. Med. Recorder, Phila. 1: 64-80, 587-600
(1818) ; Jour. de Physiol, exper. et pathologique (Magendie’s), Paris.
7: 334-384 (1827).

— . 1816 (?) On the mineralogy of Java. Batav. Genootsch. Verhandl.
8: 141-173.

. 1816 (?) Essay on the geography, mineralogy, and botany of the

western portion of the territory of the native princes of Java. Batav.

Genootsch. Verhandl. 8: 175-312.

. 1822. Systematic arrangement and descriptions of birds from the

island of Java. (Read April 18, 1820.) Linn. Soc. Trans. 13: 133-200. Oken,

Isis, 1825, col. 1053-1087.

. 1824. Zoological researches in Java, and the neighbouring island. Lon-

don.

. 1824-25. Description of the Riman-Dahan of the inhabitants of Sumatra,

a new species of Felis (F. macrocelis) discovered in the forests of Bencoolen

by Sir Stamford Raffles. Zool. Jour. 1: 542-554; Ferussac, Bull. Sci. Nat.

6: 400-402 (1825) ; Oken, Isis, 1830, col. 825-827.

. 1825. Description of the Helarctos Euryspilus, exhibiting in the Bear

from the island of Borneo the type of a sub-genus of Ursus. Zool. Jour.

2: 221-234 (1826); Ferussac, Bull. Sci. Nat. 6: 399-400 (1825) ; Oken,

Isis, 1830, col. 1023-1027.

. 1827. Notice of a species of Ursus (U. isabellinus) from Nepal. (Read

June 20, 1826.) Linn. Soc. Trans. 15 : 332-334.

. 1827-28. Notice of two new species of Vespertilionidae found in Cuba

(Molossus velox, Phyllostoma Jamaicense). Zool. Jour. 3 : 236-240.

. 1828-29. Notice of a new species of Mustela (M. Hardwickii) found

in India by Major-General Thomas Hardwicke. Zool. Jour. 4: 238-240;

Ferussac, Bull. Sci. Nat. 20: 322 (1830).

. 1828-29. Part I (-II) of a descriptive catalogue of the lepidopterous

insects contained in the Museum of the East India Company ... with...

observations on the general arrangement of this order of insects. London.

. 1829-30. Descriptions of several oriental lepidopterous insects. Zool.
Jour. 5: 65-70.

. 1831. Observations on two species of bats, from Madras (Megaderma
lyra), one of them new (Nycticejus Heathii), presented by Mr. Heath.
Zool. Soc. Proc. 1: 113-114.

. 1832. On the specific distinction of Viverra Rasse, Horsf. and Viverra
Indica, Geoffr. Zool. Soc. Proc. 2: 22-23. ‘

. 1839. A list of Mammalia and birds collected in Assam.by J. McClel-
land. Zool. Soc. Proc. 7: 146-147; Ann. Nat. Hist. 6: 366-374, 450-461
(1841) ; Oken, Isis, 1846, col. 631-634.

. 1848. Report on the island of Banka. Jour. Ind. Archipel. 3: 373-427,
705-724, 779-819; Silliman, Jour. 7: 86-101 (1849).

. 1849. Brief notice of several Mammalia and birds discovered by B. H.
Hodgson, Esq., in Upper India. Ann. Nat. Hist. 3 : 202-203.

. 1851. A catalogue of the Mammalia in the Museum of the Hon. East -
India Company. London.

. 1855. Brief notices of several new or little known species of Mammalia,
lately discovered and collected in Nepal, by Brian Houghton Hodgson, Esq.
Ann. Nat. Hist. 16: 101-104.

. 1856. Catalogue of a collection of Mammalia from Nepal, Sikkim and
Tibet, presented to the H. E. I. C. by Mr. Hodgson in 1853. Zool. Soc. Proc.
24: 393-406.

, J. J. Bennett and R. Brown. 1838-1852. Plantae Javanicae rariores,
descriptae iconibusque illustratae, quas in insula Java, amis 1802-1818, legit
et investigavit T. Horsfield; e siccis descriptiones et characteres plurimarum
elaboravit J. J. Bennett; observationes, structuram et affinitales praesertim
respicientes, passim adjecit R. Brown. London.

, et al. 1826-35. Illustrations of ornithology by Sir W. Jardine, Bart.,
and P. J. Selby, with the cooperation of J. E. Bicheno, J. G. Children, T.
Hardwicke, T. Horsfield, R. Jameson, Sir. T. S. Raffles, N. A. Vigors.
Vol. 1, 2. Edinburgh.

, and F. Moore. 1854. A catalogue of the birds in the Museum of the
Hon. East India Company. By Thomas Horsfield assisted by F. Moore.
London.

, and F. Moore. 1857. A catalogue of the lepidopterous insects in the
Museum of the Hon. East India Company. By T. Horsfield and F. Moore.
London.

, and N. A. Vigors. 1827. A description of the Australian birds in the
collection of the Linnean Society, with an attempt at arranging them accord-
ing to their natural affinities. (Read June 21, 1825 and January 17, 1826.)
Linn. Soc. Trans. 15: 170-321 ; Oken, Isis, 23, col. 258-312 (1830).

and . 1827-28. Notice of a new genus of Mammalia (Gymnura), |
found in Sumatra by Sir T. Stamford Raffles. Zool. Jour. 3: 246-249;
Ferussac, Bull. Sci. Nat. 18: 443-444 (1829) ; Oken, Isis, 1830, col. 1168-
1169. .
and . 1827-28. Descriptions of two new species of the genus Felis
in the collection of the Zoological Society (F. planiceps and F. Temmincki).
Zool. Jour. 3: 449-451.

and 1828-29. Observations on some of the Mammalia contained
in the Museum of the Zoological Society. Zool. Jour. 4: 105-113, 380-384;
Ferussac, Bull. Sci. Nat. 20: 321.

Macleay, W. S. 1825. Annulosa Javanica; or an attempt to illustrate the natural
affinities and analogies of the insects collected in Java by T. Horsfield, etc.
No. 1. London.

Notes on the Flora of Arizona

LyMAN BENSON

In this article the following topics are discussed: (1) A New
Haplophyton from the Southwest; (2) Triodia eragrostoides in
Arizona; (3) The California Poppy in Arizona.

1. A New Haplophyton from the Southwest

Dr. D. M. Crooks, head of the division of drug and related
plants of the Bureau of Plant Industry, Washington, D. C., pointed
out to the writer a difference in appearance of the Arizona plants
of Haplophyton cimicidum from figures of the same species grown
in Mexico. Investigation of the characters of specimens obtained
from the United States National Herbarium and in the University
of Arizona Herbarium has resulted in the following segregation:

HAPLOPHYTON CIMICIDUM A. DC. var. Crooksii L.
Benson, var. nov. Leaves lanceolate, 15-27 or rarely 32 mm. long,
4-8 or 10 mm. broad; seeds 6-7.5 mm. long, somewhat grooved
and ridged, commonly with part of the surface with broad papillae
resembling pebble-grained leather. Foliis lanceolatis, 15-27 mm.
rariter 32 mm. longis, 4-8 mm. rariter 10 mm. latis; seminis 6—7.5
mm. longis, striatis vel partim papillatis. Southeastern Arizona to
Western Texas; southward into Northern Mexico. Type collec-
tion: “Prison Road,” Santa Catalina Mountains, Pima County,
Arizona, D. M. Crooks & Robert A. Darrow, Dec. 27, 1939. Type
mounted on three sheets in the Herbarium of the University of
Arizona.

The corresponding characters of typical Haplophyton cimicidum
are as follows: Leaves ovate-attenuate, 35-45 mm. long, 14-22 mm.
broad; seeds 8-10 mm. long, deeply grooved and ridged. The
species is common in southern and. central Mexico, and it occurs
as far northward and westward as Guaymas, Sonora (Palmer in

1887, U. S.). Specimens of the variety with leaves large enough to
be considered almost but not clearly transitional are the following:
Baboquivari Mountains, Arizona, Peebles, Harrison & Kearney
2795, U. S.; Rio de los vueltos, Mexico (state not given), Lieb-
mann 11993, U. S.; Eulalia Plains, Chihuahua, Wilkinson in 1885,
Oo Sc

Haplophyton cimicidum is known as “hierba de la cucaracha”
or cockroach plant, and the vegetative parts contain an insecticide
used with cornmeal to kill cockroaches.

2. Triodia eragrostoides in Arizona

Triodia eragrostoides Vasey & Scribn. is one of many species
growing in northern Mexico, which occur in Arizona and Texas
but not in the intervening area in New Mexico. It has not been
reported heretofore for Arizona. Mesquites along a small wash at
the Barbeque Area of the Colossal Cave State Park, Pima County,
Arizona, L. Benson 9174, Sept. 28, 1938, L. Benson 9801, Oct. 9,
1939. Range, cf. A. S. Hitchcock, Manual of the Grasses of the
United) States 213511935, ~Hlorida Keys, Wexas, and) northern
Mexico; Cuba,” or, cf. W. J. Beal, Grasses of North America
2: 465. 1896, “Florida, Texas, and Mexico.”

3. The California Poppy in Arizona

The California poppy, Eschscholtzia californica Cham. presents
a classification problem to the systematic botanist, wherever he may
find it, and it is not surprising that the plant occurring on the desert
plains and hills in central and southern Arizona is unusual in some
respects. It is difficult to discover enough characters in the California
poppy to match the hundred or so specific names proposed by
Greene, Pittonia 5: 205-293. 1905, but the species is variable in
California. The annual form growing in Arizona is readily matched
by some California plants, but it does not agree in some characters
with the bulk of plants in that state. The torus rim is either not
present or reduced to a ring not more than 2 mm. broad, the stems .
have a tendency to be scapose, and most years the flowers are smaller
and paler. However, the excellent rainy spring of 1941 afforded an
opportunity for study of the Arizona plant under conditions ap-
proximating those in various parts of California. According to the
field observations of the writer, there is no reason to provide the

Arizona plant with a name other than Eschscholtzia californica, and
specific names such as E. mexicana Greene, E. aliena Greene, E.
Jones Greene, FE. arizonica Greene, and F. paupercula Greene
(cf. Greene loc. cit. pp. 260-263) are merely metanyms.

It is noteworthy that flower color is more variable than in the
California forms of the species. In the poppy fields near Tucson
colors included orange, yellow with orange center, white with yellow
center, white, and numerous variations in color intensity within the
major groups. Similar color-types occur in California, but those
other than orange or orange-yellow are uncommon in the spring-
time, while in Arizona they are remarkably prominent.

DEPARTMENT OF BOTANY
UNIVERSITY OF ARIZONA
Tucson, ARIZONA

The Names of Cornus

H. W. Rickert

So early as 1833 Lindley, in founding his genus Benthamia
(Bot. Reg. 19: 1579 et seq.), remarked “We do not understand
upon what principle this very distinct genus has been combined
with Cornus, from which it differs essentially both in flower and
fruit. Whether or not C. florida, which agrees with it in habit, is
also a species of Benthamia, our materials do not enable us to de-
termine.” In 1828 Rafinesque (Med. Bot. 132) had distinguished
C. florida as section Cynoxylon, which in 1838 he elevated to ge-
neric rank (Alsog. Am. 59). This early tendency to divide the
genus has continued, with varying success, until modern times.
For instance, Moldenke (Rev. Sudam. Bot. 6: 177. 1940) says:
“There is certainly no doubt in my mind that the genus Cornus as
regarded by many botanists today is actually an aggregate of several
distinct generic elements. The true genus Cornus is typified by
Cornus mas L. and contains the so-called Cornelian-cherries. The
cornels or osiers represent the genus Svida, the bunchberries repre-
sent the genus Chamaepericylmenum, the American flowering-
dogwoods represent the genus Benthamudia, and the Asiatic
flowering-dogwoods with their coalesced fruit represent the genus
Benthamia.”

Aside from the taxonomic question here involved, the nomen-
clature of these segregates repays scrutiny. To begin at the begin-
ning, when Lindley founded Benthamia (1.c.) he said of the name:
“The Benthamia of Achille Richard being the same as Herminium,
we have great pleasure in availing ourselves of the present oppor-
tunity of naming this very distinct genus in compliment to our
highly valued friend George Bentham, Esq.’ The sentiment did
him honor, but the result is inconformable with our rules of nomen-
clature, Benthamia Richard, an orchid, having been validly pub-
lished in Mém. Soc. Hist. Nat. Paris 4: 37 (1838) .*

Benthamidia Spach (Hist. Vég. Phan. 8: 106. 1839) is ante-
dated by Cynoxylon Raf. (Lc.). I cannot agree with Farwell
(Rhodora 34: 29-30. 1932) that Cynoxylon was not intended for
generic rank. It is true that Rafinesque did not make combinations
under his new name; true also that he did not always make his
intentions plain. But to unriddle Rafiinesque’s intentions and, above
all, to expect consistency in his writings, are beyond the powers
and the prerogatives of a scientist. Speaking of his segregates as
“G. or subgenera,” he lists “255. Subg. Mesomera Raf. 256. subg.
Kraniopsis Raf. 257. EUKRANIA Raf. 258. CyNoxyton Raf. 259.
BENTHAMIA Lind.” (lc. 58-59). Each is briefly characterized.
He goes on to “mention all the true Cornus,” the species included
in the first two groups.

Eukrania Raf. (1.c.) included as “types’”® C. mascula, C. cana-
densis, C. suecica, Of this odd assortment C. mas L. (“C. mascula’)
has been designated as the type of Cornus L. The change in the
circumscription of Eukrania by the removal from it of C. mas (or,
to put it differently, the division of the genus) does not invalidate
the name, which must be retained if the “bunchberries” are to be
treated as a genus. Eukrania Raf. of course antedates Chamae-
pericylmenum Graebner (Asch. & Graebner. FI. Nordostdeuts.

1Tt is interesting also to note a previous abortive attempt by Lindley to
name a genus after Bentham (Nat. Syst. 241. 1830, nomen nudum), appar- -
ently a genus of Boraginaceae and according to A. de Candolle (Prodr.
10: 118. 1846) used on labels in the garden of the Horticultural Society.

* Rydberg wrote (Bull. Torrey Club 33: 147. 1906) that Rafinesque made
C. mas “the type” of the genus. In 1839 Rafinesque was far from designating
nomenclatural types in the modern sense. Actually he named three species
as “types,” by which he must have meant “typical.”

Flachl. 539. 1898), and Cornella Rydb. (Bull. Torrey Club 33:
147. 1906).

Svida is derived from a Czech word for dogwood. Opiz
(Seznam 94. 1852) made it a genus-name and referred to it
C. sanguinea L., the common European shrub called dogwood
in England,* and C. alba L., related to our C. stolonifera Michx. ;
but failed to describe it. Indeed, we can infer his intention to divide
Cornus only from the existence of C. mas on page 33 of his flora.
Such a procedure, though legitimate at the time, is contrary to our
present rules. Svida was first validly published by Small in 1903
Gal, S12, We Se S96).

There are those who say that such a disturbance of the dead
bones of nomenclature can be prompted only by the disturber’s
desire to see his name after new names and combinations. Per-
haps I should grasp the opportunity to give the Asiatic flowering
dogwoods a legitimate name and to make new combinations under
Eukrania Raf. emend. But botanical bibliography is the servant
of taxonomy ; this catalogue of oversights is only incidental to the
revaluation of the groups. The point is that a consideration of the
genus Cornus over its entire range renders its division far less easy.

Cornus Volkensii Harms (in Engler, Pflanzenw. Ost-Afr. C:
301. 1895), the only known species in Africa, has a paniculate
inflorescence much like that of the European C. sanguinea but
enclosed in four early deciduous bracts like those characteristic
of C. mas (southeastern Europe and western Asia). The drupe
is ellipsoidal as in C. mas but dark-colored as in C. sanguinea. It
fits neatly as an intermediate between the sections which include
these species. C. disciflora Moc. & Sessé (ex DC. Prodr. 4: 273.
1830) of Mexico has the “capitate” inflorescence (a reduced pani-
cle) of our C. florida but its four bracts are small and early decidu-
ous as in C. mas and C. Volkensu,; its drupe is ellipsoidal and
dark-colored. There seems to be a tendency toward dioecism (char-
acteristic of several genera of Cornaceae) in both C. Volkensu and
C. disciflora. The concrescence of the fruit characteristic of the
Asiatic C. Kousa seems hardly to warrant generic segregation,
especially since it is approached by C. Nuttalli: of our west coast.

3 Not, of course, an “osier,” though C. stolonifera is often called the “red
osier.” Osiers are properly willows; the name has sometimes been used for
other withe-like shrubs similarly used in Europe for constructing wattles.

As for Eukrania, it bases its claims to recognition on its “herba-
ceous’”’ habit and the presence of a small dorsal horn on the petals.*
In several characters it is intermediate between C. florida and the
ebracteate dogwoods.

I do not know what we are to understand by such expressions
as “an aggregate of generic elements.” They may signify that
Cornus (sensu lato) is polyphyletic, distinct genera having been
merged; or that an original stock has become diversified. The
latter seems more plausible. Certainly in our ignorance of the
history and cytogenetics of the group the burden of proof must
fall on him who expounds a polyphyletic origin; present judgment
seems premature. If they are really as closely related as they seem
to be, I see nothing to be gained by segregating in distinct genera
the seven (not five) recognizable sections of Cornus.

New York BotTanicAL GARDEN
New York, N. Y.

Phyllanthus nummulariaefolius Poir. in the United States

LEoN CRoIzAT

About five years ago correspondents in Brazil and the Panama
Canal Zone sent me seeds of an undetermined species of Phyllan-
thus which they described as a polymorphous and aggressive weed.
I planted these seeds in a hothouse, grew out of them a sizable
crop of specimens and satisfied myself that P. nummulariaefolius
Poir. was the entity that had been collected. This plant has proved
to be as aggressive and as polymorphous in the hothouse as it has
been reported to be in nature, and I must now carefully eradicate
it several times a year. The size of the specimens varies from a few
inches tall, when the plants happen to grow on a dry bench, to about
three feet for material favored by good conditions of soil and
temperature.

Pressed specimens of the same plant have also reached me from
Argentina, Brazil, Panama and the French West Indies, showing
that it is widespread in every one of the tropical American coun-

# Rydberg (1.c.) said on the sepals; surely an error.

tries bordering upon the Atlantic Ocean. In no case has the mate-

rial thus sent proved to be correctly determined, being usually
mislabelled as P. Niruri L. or P. lathyroides H.B.K. These mis-
determinations are not always excusable because P. nummulariae-
folius not only manifestly differs from both those species and their
nearest allies, but represents in the American flora a type of vege-
tation that has no immediate relatives. Its affinities are African and
Asiatic.

Léandri, who has contributed several specimens to our herba-
rium and has extensively collected this weed in its endemic range,
that is, Madagascar and the adjacent islands, is the author of a ©
critical study (in Lecomte Not. Syst. 7[4] :168-169, 171-172, 1939).
Here, he stresses the impossibility of using the relative size of the
leaf and the length of the fruiting pedicel to separate, even tri-
nomially, the many polymorphous aspects of the species. Léandri
treats P. tenellus Roxb. as a synonym of P. nuimmulariaefolius, a
disposition which is fully justified by the material of the latter
which I have seen in the Kew Herbarium, part of which at least
was seen by Hooker when preparing the classic illustration of
P. tenellus (in Hook. Icon. 16: Pl. 1569. 1887). It is quite evident
that P. minor Fawcett (in Jour. Bot. 57:65. 1919) is a synonym
of P. nummutlariaefolius, from which Fawcett attempts to sepa-
rate it on the basis of minor vegetative characters. An isotype of
P. minor in the herbarium of the N. Y. Botanical Garden, Harris
12123, fully matches specimens of P. nummulariaefolius such as
grow in moist and shady situations in a hothouse. I believe, more-
over, that Lanjouw is justified in suggesting (in Rec. Trav. Bot.
Neerl. 31:452. 1934) that P. corcovadensis Muell. Arg. is a
synonym of P. nummulariaefolius and an African weed introduced
into America. I have not yet seen authentic material of Mueller’s
species, but its description and illustration (in Martius Fl. Bras.
11[2] :30, Pl. 6 ii. 1873) apply to no other plant better than to
Poiret’s Phyllanthus.

Rio de Janeiro apparently was the original point of introduc-
tion of this noxious weed into America, having been brought there
probably by ships sailing in colonial times between Mauritius and
Brazil. I may note that this is not the only record of an introduc-
tion of the kind. Euphorbia spathulata Lam., the holotype of which
I have seen, is supposed to be endemic to the Plata regions of

Argentina, but is altogether alien to the native flora of South
America, and it so well matches FE. dictyosperma Fisch. & Mey. and
the minor segregates in its vicinity as to suggest that the alleged
Argentina endemic is but the North American weed, introduced in
the regions of the Plata before 1780. It is characteristic that Norton
lists (in Rept. Mo. Bot. Gard. 11:104. 1900) a Moyer specimen
from Montevideo under Euphorbia arkansana Engel. & Gray var.
Missouriensis.

In view of the widespread range and of the aggressiveness of
P. nummutlariaefolius | have been looking forward to finding it
recorded within the continei:tal limits of the United States some-
where along the coast between Texas and the Carolinas. My antici-
pations have been only very recently fulfilled by the finding of two
specimens in the herbarium of the N. Y. Botanical Garden, namely :
Moldenke 151, Orlando, Fla., 1929, and Rapp 3, Sanford, Fla.,
1932, which unmistakably belong to this species. So far, I have
seen no other specimens collected in the United States and accept,
consequently, Moldenke 151 as the first record of P. nummula-
riaefolius for the flora of the United States, exclusive of its ter-
ritories and dependencies.

Phyllanthus lathyroides is reported by J. K. Small for Florida
(Man. Southeast. Fl. 778. 1933), but he does not mention either
P. nummulariaefolius or its synonyms, P. tenellus and P. corcova-
densis. Since the Moldenke and the Rapp records have been origi-
nally misdetermined as P. lathyroides, and the former has certainly
been seen by Small when at work on the Manual, I suspect that
the record of P. lathyroides in Small’s work is based upon a mis-
determination. I have not seen material of P. lathyroides from
Florida, but this species.is likely to have been introduced there,
and Small may thus have seen authentic specimens which are now
not preserved in the herbarium of the N. Y. Botanical Garden. He,
at any rate, failed to record P. nummulariaefolius.

Taxonomists who are interested in learning how to distinguish
P. lathyroides from P. nummulariaefolius should study actual speci-
mens rather than rely upon the compilations and the colorless
descriptions so frequently found in the literature. The two species
are quite distinct and excellent material of both is preserved in
the herbarium of the N. Y. Botanical Garden. The following speci-
mens represent P. lathyroides in that herbarium: (1) Britton,

Britton & Brown 6995, Portorico; (2) Britton & Boynton 8201,
Portorico; (3) Duss 47, Martinique, French W. I., and are true to
the isotype which I have seen in the Parisian Museum.

Phyllanthus nummulariaefolius (=P. tenellus Roxb. ; P. corco-
vadensis Muell. Arg., syn. nov.; P. minor Fawc., syn. nov.) is
represented by the following collections: (1) Ball s.n., 1882, Tijuca,
Rio de Janeiro, Brazil; (2) Duss 2442-5557 [duplicate sheet],
Guadeloupe, French W. I.; (3) Harris 12157, 12208, 12123 [three
sheets, including isotype of P. minor], Jamaica.

The best characters of identification of P. nummulariaefolius
from P. lathyroides and the species or forms in the latter’s vicinity
(e.g., P. diffusus Kl., well represented by: J. S. De La Cruz 3662,
British Guiana, in the herbarium of the N. Y. Botanical Garden)
are the following: (a) Shape of the leaf. In P. nummulariaefolius
the leaf, regardless of its size, is more or less gradually narrowed
from the center towards the extremities, being ovate to obovate.
In P. lathyroides and P. diffusus the leaf is essentially elliptic,
with the sides tending to run more or less parallel. (b) Length of
the pedicel. In P. nummulariaefolius the pedicel, especially that
of a fruiting flower, is subcapillary but stiffly produced, always
manifestly elongate. In P. latnyroides and P. diffusus the pedicel is
much shorter. In P. miruri the pedicel is very short, so that the
female flower can here be described as subsessile. (c) Size of the
lobes of the calyx of the female flower. In P. nummulariaefolius the
lobes are small, narrowly triangular-acuminate, showing like a
minute “star” at the tip of the pedicel. In P. lathyroides the lobes
are definitely large and subpetaloid. In P. diffusus and P. Niruri
the lobes are much smaller than in P. lathyroides and thus tend to
approach the size if not the shape of those of P. nummutlariaefolius,
but the length of the pedicel is much shorter, as noticed above.

The seed furnishes good characters of determination in Phyl-
lanthus, but only mature seeds can be usefully compared for critical
identifications and it is unfortunate that there are all too few speci-
mens in herbaria which have a complement of seeds fit to be used.
The vegetative characters listed above will be found adequate, I
believe, at least for provisional determinations.

THE ARNOLD ARBORETUM
Jamaica Prain, Mass.

BOOK REVIEWS

A New Text for College Botany

The Plant-World, A Text in College Botany. By Harry J. Fuller.
Henry Holt and Co. 1941. Pp. 592. $3.25.

Another excellent text has been added to the ones planned
for a first course in college botany. With so many excellent texts
already a new one should justify itself by some difference in its
approach to the subject, in the aspects of the science stressed, or
in the special group of students for which it is planned. In the
preface the present text explains that it is for “students who are
registered in elementary botany courses principally because of the
cultural and general educational value of the subject” and who
presumably will take no other courses in biological subjects. With
this in mind the author has chosen and arranged the subject
matter with the idea of arousing the interest of the students at
the start by associating the study of plants with their everyday
experiences. The primary objective given is “the presentation of
the fundamental features of structures, physiological activities, and
reproduction of flowering plants.” Considerably more than half
the book is devoted to this main objective. Of several secondary
objectives the presenting of a generalized account of plant evolu-
tion is given last, with the suggestion that the section of the book
treating it and plant ecology may be omitted. Thus many students
using the text will undoubtedly finish the course without getting
even the brief description of evolution given in the text. The struc-
ture and classification of plants below the Spermatophytes is given
very briefly, as is heredity and plant breeding.

The short introductory chapters on the history of botanical
study and on the nature and origin of life are well done and should
stimulate interest at the start. Conforming to the announced
objectives the classification of plants is taken up only briefly, using
as “a pedagogical concession” the old grouping into Thallophytes,
Bryophytes, Pteridophytes and Spermatophytes; though an out-
line of a more modern system of classification is given in an
appendix.

Illustrations are many and excellent, the drawings, photographs
and photomicrographs are good and well reproduced and are
chosen to really illustrate the text. The frontispiece is a beautiful

colored picture of a Cattleya, but, as is often the case in text books,
it is merely a pretty picture not in any way important to the book.

As in nearly all college science texts—and the same is true in
only slighly less degree of high school texts—the student will meet
here nearly as many new terms as he will new words in the first
year of a foreign language. The glossary gives nearly 600 tech-
nical terms, most of which will be new to the student, while others
(such as xeromorphic, polyploidy, photophobic) used in the text
are not given in the glossary. It may be difficult to draw the line
as to which scientific terms should be included and which omitted
in a book of this kind, but for students most of whom will take |
no further botany it seems unreasonable to require the learning
of scores of words used but once in the text—and there with an
explanation—and which they may never in their lives meet again.

There is nothing in the text to suggest laboratory or field
work, nor references to further reading. Each chapter is followed
by a concise summary, which correctly used, will be a definite help
in mastering and organizing the facts given. The language
throughout is clear and easily understood, so that the book may
be read by a beginner with pleasure. It should satisfactorily fulfill
the author’s objective for the course. It will be a valuable text
wherever a cultural course in botany, not to be followed by more
advanced work, is given. The reviewer hopes that whenever the
text is used part [V—‘“‘The Distribution of Plants in Time and
Space” will not be omitted.

Georce T. HASTINGS

The Advance of the Fungi

The Advance of the Fungi. By E. C. Large. Henry Holt and Co., New
York. 1940. Pp. 488. $4.

Under the above title one would naturally expect to find a
discussion of either the phylogeny of fungi in general or a myco-
logical treatise. A glance at the chapter headings may have a rather
discouraging effect on the young plant pathologist, for here he
would find little information about individual plant diseases, which
might be expected in a work on plant pathology. Nevertheless, the
author deals primarily with plant-pathological problems, availing
himself on every occasion of setting forth some of his philosophical
or sociological ideas.

The two opening chapters on potato murrain and the famine
in Ireland contain little not already familiar to the mature plant
pathologist. The young student might expect to find at the close
of these chapters something on the modern methods of the control
of the disease. We might also expect the author to take this oppor-
tunity to answer some of those who have been criticizing the
scientist because of the way his discoveries have been utilized in
the construction of the deadly weapons of modern warfare. He
could, in accord with Dr. Blakeslee’s recent address as retiring
president of the A. A. A. S., have pointed out the great contribu-
tions for good made by scientists who have shown how, for exam-
ple, potato blight can be easily controlled so that famines in Ireland
are no longer necessary or probable. In later chapters on Bordeaux
mixture and “New Sprays for Old” methods are given for con-
- trolling the blight.

When one considers the author’s sociological viewpoints he finds
an excuse for a good discussion of Phylloxera even though aphids
are not very closely related to the fungi! This chapter on Phyl-
loxera would naturally be the last place one would look to find an
account of Craigie’s discovery of the functioning of the spermatia
of wheat rust, which would naturally be included in the chapter
on the “Barberry and the Wheat.” Craigie’s work, however, is
also mentioned in the chapter “Towards Immunity” where the
origin of many of the new biologic races is properly attributed to
hybridization in the wheat rust.

A chapter on degeneration and virus diseases is included, no
doubt because viruses as well as fungi cause disease. Here the
author has briefly yet effectively given us the latest information
on this type of disease.

On the whole one cannot help enjoying a leisurely reading of
various chapters because the historical accounts of certain of our
most destructive plant diseases are enlivened with ideas on human
relations well worth pondering.

B. O. DopcE
New York BoTANICAL GARDEN

Zyl

An Unusually Good Book

Hunger Signs in Crops. A symposium written by a group of fifteen spe-
cialists in agronomy, horticulture, plant nutrition, and plant diseases. Pub-
lished by the American Society of Agronomy and the National Fertilizer
Association. Judd and Detweiler, Inc., Washington, D. C. 1941. Pp. 340. $2.50

Hunger Signs in Crops gives in a very practical manner the
symptoms that develop in growing crops when they lack needed
mineral elements. The book is timely, for nutritional experts inform
us that our diets are woefully lacking in vitamins, proteins and
minerals. When plants lack minerals they cannot grow normally,
and man and animals that feed upon these plants do not obtain the
essential food elements.

The seventy-nine color plates in the book are well chosen and
illustrate clearly the many points emphasized throughout the vol-
ume. As an example, the picture of a grapefruit with aborted seed
and gum pockets in its axis clearly shows boron deficiency. The
normal fruit in section is shown for comparison. In addition to the
colored plates there are ninety-five halftones that vividly show the
results of mineral deficiencies in the plants. The plants discussed
are the ones we deal with in our daily life. The pictures illustrate
the poor vegetables and fruits that we often purchase unwittingly
from the store.

The opening chapter deals with general considerations but fol-
lows with a discussion of tobacco, corn and small grains, potato,
cotton, vegetables or truck crops, deciduous fruit, legumes and citrus
fruits.

The book was designed to be non-technical so as to increase its
usefulness. The material was planned for county agents, agricultural
teachers, progressive farmers and a source book for libraries and
scientists. The clear pictures show at a glance what is wrong with a
plant. Thirty minutes spent in the projection of the splendid plates
will teach a student more about mineral deficiencies than ten hours
of didactic work. Botanists and all lovers of nature cannot afford to
ignore this book if they wish to be classified among the well in-
formed.

As one turns the pages of the book one is confronted with the
need of the following fertilizers in the soil: nitrogen, potassium,
phosphorus, sulfur, magnesium, calcium, iron, manganese, boron,
zine and copper. When these elements are lacking, we have the ready

picture which shows the deficiency and the loss of yields that one
may expect. As a defense measure crops must be of high quality,
and proper plant nutrition is absolutely necessary if we are to pro-
duce in abundance.

Sales of this book have been unusually high which testifies to its
real worth. Credit, however, must be given to its sponsors who con-
tributed freely of their time and to the Soil Improvement Commit-
tee of the National Fertilizer Association who agreed to be respon-
sible for the sale of enough copies so that the price of the book would
be within reach of all.

ForDHAM UNIVERSITY Wo. J. BoNISTEEL.

PILED TRIPS OF DAE Cuus

Trip OF SEPTEMBER 28, 1941, TO LAKE BEAR Swamp (LAKE
OwASSA) AND SPRINGDALE, N. J.

This was a joint outing with the American Fern Society. Our
first find was made before reaching the swamp. Among the revege-
tating species in a long abandoned field at the edge of the swamp
we found the two gentians typical of north Jersey, Gentiana quin-
quefolia and G. Andrewsti. Two species of Botrychium were taken
here also. In rapid succession as we entered the swamp the Massa-
chusetts fern, and the two chain ferns were encountered. All of
the species commonly to be expected in this habitat were found.
Our trip had been prompted by the leader’s interest in a press re-
port that “mining” operations were in progress in the vicinity. It
seems that a so-called “peat’’ is obtained from the root masses
(tussocks) of Osmunda. No evidence of such activity was encoun-
tered though Osmunda was plentiful. This is a large swamp and
we did not cover it all, though the difficulty of crossing a sector
of Rhododendron thicket convinced most of the party that they
had travelled miles. The reward here was a good feed of wild grapes
in their prime. Before leaving the parking place many of the group
were successful in finding Isoetes along the shore of Lake Owassa.

After lunch we returned to Newton and the leader obtained
permission from Mr. Whittingham to cross his pasture to the well-
known Springdale swamp region. Many previous visits to this area
have been made. Clinton’s and Goldie’s ferns are abundant in parts

of the swamp as well as numerous other species of Dryopteris.
During the past forty years many hybrid forms have been dis-
covered by the members of the two clubs. One such colony of
Goldiana « Marginalis was visited. This colony was first reported
by Philip Dowell. At this time it was found to contain several
plants, generally in good condition despite the dry season. Two
plants of hart’s tongue fern were planted here by the Fern Society
some years ago. Mr. Leon Bowen had reported them in good con-
dition last winter. We found one plant to have nine good-sized
leaves, eight of them fertile. No signs of reproduction were to be
seen. The other plant was in poor condition so it was reset in the
hope of finding more congenial surroundings. The leader pointed
out that the soil and rock conditions of the native habitat in central
New York are similar but the slopes are higher and cooler there.
No visit to the Springdale swamp would be complete without in-
cluding the Big Spring. There is a large colony of the common
water buttercup here, Ranunculus delphinifolius. It was in flower
at this date. On other trips we have collected it in flower as early
as May 15, indicating a possible flowering period of nearly five
months. Attendance: about 30. Leader: R. C. Benedict.
JoHN A. SMALL

Trip oF JUNE 21-JuLy 5—Eastern New ENGLAND Tour

This trip of some 1800 miles was held substantially as announced
in the field schedule. The hotel selected on Mt. Monadnock was the
Half Way House which we found completely adequate. Plants of
the Canadian and sub-alpine zones were seen on Mt. Monadnock,
some of them in great beauty and abundance. Forestry practices
and the destruction caused by the hurricane in 1938 were seen at
the Caroline A. Fox Research and Demonstration Forest. Both of
these walks were led by Dr. Henry I. Baldwin. Dr. Albion Hodgdon
gave us some good trips in the Durham vicinity, stressing the
behavior of plants at the end of their range. A northern bog, cedar
swamp, and various upland situations were examined.

Mr. Arthur H. Norton of the Portland Society of Natural
History, assisted by the botanists of the University of Maine, gave
us a tour of York County in southwestern Maine. Sand barrens,
bog lake, seashore, salt marsh, and fresh marsh were included.
Intermediate stops were made at stations for particular plants of

local occurrence. We climbed Mt. Agamenticus (alt. 673 ft.) for
a grand view of the surrounding country. This is the high point
of York County and is of local importance in being near the shore,
forming a landmark in the monotonous coastline as viewed from
the sea. Of course it figures in local nautical yarns. To us it brought
Selaginella rupestris, Juniperus communis, and a dwarf species of
Amelanchier, in addition to the more common species of the maple-
oak forest. An old friend Arctostaphylos Uva-ursi was found here
growing over the exposed granite.

Mt. Washington was a high point in many ways. Both Pinkham
Notch Camp and Glen House were delightful though quite different.
We had two splendid days. The two endemics, Geum Peckii and
Houstonia caerulea var. Faxonorum were abundant and in full
bloom. Dr. Baldwin arranged a fine symposium in the Alpine
Garden with speakers who knew the region from first-hand re-
search. These included Dr. Richard Goldthwaite on geology, Dr.
R. F. Griggs on ecology, Mr. Norton on birds, Dr. C. F. Jackson
on mammals, Dr. S. K. Harris on plants, a representative of the
Mt. Washington Observatory on climate, and a member of the
Forest Service on management policy of the White Mt. National
Forest.

Two equally spectacular days awaited us at Mt. Katahdin. A
five-mile hike to and from the northern terminus of the Appala-
chian Trail. A climb down and up the Chimney. Overnight in lean-
tos on bough beds. Meals by a Maine guide or at a Maine sporting
camp. Crossing the summit in clouds. All these conspired to enhance
our pleasure in seeing the many species of alpine plants to which
Dr. F. H. Steinmetz led us. The heat of the sun on the mountain
table-land, the cold of the mild storm, snow in protected ravines,
high winds, steep slides, cliffs, dry exposed rock, springs, and
Chimney Pond were some of the varied habitats that we examined.
The response of forest species to altitude and these other factors
was carefully noted by Dr. Pierre Dansereau of the Montreal
Botanical Garden.

The trip through eastern Maine was no less outstanding for
Dr. Steinmetz went to unending pains to show us unique habitats
and particular species of plants. Streams, the stony coastal head-
lands, the raised bogs or high moors, and the blueberry barrens
were accompanied by most interesting elaboration of their environ-

ment and floristics. Good lodgings and intriguing meals ranging
from a picnic with “makings” obtained at a four corners store
(which had been in business for over 100 years) to a complete
Maine shore dinner kept us in trim for the long days collecting and
the short evenings (nights) for pressing.

Finally a day in Acadia National Park with Maurice Sullivan,
Park Naturalist, brought our tour to a close. Species have not been
mentioned in this report because of the vast number that were of
interest and the limitations of space. Lists from characteristic
habitats and local stations of botanical significance have been pre-
viously recorded by others and are available. A possible extension
of range in the discovery of Iris setosa at Jonesport by Dr. Jacques
Rousseau of the University of Montreal is our only chance of
contributing to botanical science. Daily attendance fluctuated from
seventeen to fifty-eight. Total participation was seventy-five. A
final word of thanks to all who guided us.

Joun A. SMALL

BNO CEE DINGSEOH rit @le eis

MINUTES OF THE MEETING OF OcToBER 15, 1941

The meeting was called to order by the First Vice-President,
Dr. E. B. Matzke, at the New York Botanical Garden at 3:30 P.M.
Thirty-five members and friends were present.

In the absence of the Recording Secretary the Corresponding
Secretary read the minutes of the previous meeting. These were
adopted with correction.

It was voted that Miss Mary Gojdics, Duchesne College, Omaha, Neb.,
be unanimously elected to annual membership.

The Corresponding Secretary requested the permission of the
Club to have its name used in the press as being opposed to the
proposed amendment to the State Constitution which would permit
construction of a ski trail on Whiteface Mountain. After discus-
sion, it was moved by Dr. Camp that this permission to use the
Club’s name be granted. Dr. Kolk seconded the motion and the
Club so voted.

The scientific part of the program consisted of two discussions
illustrated by lantern slides and living specimens. The first speaker,

Dr. John D. Dwyer, spoke on “Interesting plants of Litchfield
County, Connecticut.” The speaker’s abstract follows:

A summer and fall survey of the flowering plants and ferns growing
on a 4,000-acre tract of land in Litchfield County, Connecticut, and super-
vised by the State Board of Fisheries and Game, yielded approximatetly 600
species. Since the tract surrounds Bantam Lake and includes several ponds,
opportunities for the study of aquatic vegetation were offered. Seventeen
species of Potamogeton, including seven varieties, were collected. Numbered
among these is P. bupleuroides Fernald, hitherto not reported for Connecticut
west of Windsor Locks. Special collections and study of the complex species,
Arisaema triphyllum were made. Kodachrome studies of exceptional and
attractive plants were featured.

The second speaker, Mr. Jerome Metzner, spoke on “Observa-
tions on Local Volvocales.” The speaker’s abstract follows:

The three local species of Volvox may be distinguished from each other
easily on the basis of certain differences in vegetative characteristics.
V. globator has lobate protoplasts which are connected to each other by stout
protoplasmic connections containing contractile vacuoles. V. aureus is about
one-half the size of VY. globator. Its protoplasts are not lobate and are con-
nected by very delicate strands of protoplasm. V. weismannia is approxi-
mately the same size as VY. aureus but lacks completely any protoplasmic
connections. The oospores of V. globator are large and possess stout conical
spines. The oospores of V. aureus lack spines. In V. weismannia there are
slight spiny projections from the surface of the oospore.

Our knowledge of the life cycle of the genus Volvox is incomplete since
fertilization has never been seen in any species. Preliminary studies made
at Barnard College seem to indicate a complete lack of fertilization in
V. weismannia. The oospores may be partenospores. Studies made on the
development of the juvenile colony from the oospores in V. weismannia
reveal the presence of protoplasmic connections in the early stages. This is
possibly indicative of the ancestral condition.

The meeting was adjourned at 4:35 P.M. to enjoy the refresh-
ments served by the members of the Garden staff.
Respectfully submitted,

JOHN W. THOMPSON, JR.
RECORDING SECRETARY

MINUTES OF THE MEETING OF NOVEMBER 3, 1941

The meeting was called to order by the President, Dr. J. S.
Karling, at the American Museum of Natural History at 8:15 P.M.
One hundred and eleven members and friends were present.

The minutes of the previous meeting were adopted as read.

It was voted that Dr. Flora Murray Scott, University of California,
405 Hilgard Street, West Los Angeles, Calif., be admitted by unanimous
ballot to annual membership in the Club.

The scientific part of the program consisted of a talk by Dr.
E. B. Matzke of Columbia University on “Autumn Coloration.”
‘The speaker’s abstract follows:

When the green pigments, chlorophyll a and chlorophyll b, break down
in the fall of the year, the carotene and xanthophyll, which are yellow to
reddish-orange, become evident; anthoxanthins may be pale yellow. Antho-
cyanins are responsible for the brilliant red to violet colors of certain plants ;
their formation is governed by the genetic make-up of the plant, internal
nutriment, light, temperature, available water, fixed nitrogen, and oxygen.
The final brown is caused largely by tannins.

Through New England the sugar maple is the tree most largely respon-
sible for the colors of autumn—varying from yellow to brilliant red. Its
counterpart farther south is the scarlet oak, though other species of oak
are also important. Red and purple colors are also added to the landscape
by the dogwood, sour gum, sweet gum, sassafras, and white ash. The yellows
are largely furnished by the hickories, tulip tree, and ginkgo. Black cherry,
last of our trees to turn, takes on all colors, from purple to yellow.

Among the shrubs, purples, reds, and yellows are added by the sumachs,
blueberries, barberry, and spicebush. Vines like cranberry, Virginia creeper,
and Boston ivy, add their more modest bit. In the salt marshes glasswort
is brilliant red. Beard grass paints the poorer hillsides tawny orange. Fruits,
like those of holly, bittersweet, hawthorne, and barberry, each add their touch
of red or yellow.

This display is characteristic of eastern Asia and eastern North America;
in Europe, the Danube valley and parts of Switzerland are also showy,
but to a less extent.

This final fanfare of color has no deep underlying biological significance.

The meeting was adjourned at 9:25 P.M.
Respectfully submitted,

JOIEOND WY, “WislOMWMIPSOIN, IR
RECORDING SECRETARY

MINUTES OF THE MEETING oF NoveMBER 19, 1941

The meeting was called to order by the First Vice-President,
Dr. E. B. Matzke, at 3:35 P.M. at the New York Botanical Garden.
Thirty-two members and friends were present.

In the absence of the Recording Secretary, the Corresponding
Secretary read the minutes. The minutes of the previous meeting
were adopted as read.

Mr. John T. Presley, Sacaton, Ariz., was elected by unanimous ballot
to annual membership.

The scientific program consisted of three talks. The first
speaker, Mr. Robert Hulbary, discussed “A fungus disease of
Austrian pine.” The speaker’s abstract follows:

In blighted needles of Austrian pine collected in northern Illinois in the
fall of 1938, immature stromata indicated the cause of the blight. Infected
needles were wintered out-of-doors and examined periodically. The stromata
remained quiescent through the winter but very early in the spring began to
develop and by March 1 had emerged as strongly erumpent, loaf-shaped
structures. A month and a half later, pycnidial locules were becoming dif-
ferentiated, and by May 15 conidia were being produced.

The distinctive dothideaceaceous structure of the stroma distinguished
the fungus from every described group. For it the new genus Dothistroma
is proposed.

The well-marked dothideaceous structure of the stroma and the spore
characters place the new fungus in the scolecosporous group of the Phoma-
ceae close to Hemidothis Sydow. and Septocyta Petrak.

The second speaker, Mr. John Dodd, discussed “Some reactions
to grafting in Viola.”

The third speaker, Dr. Sydney Greenfield, discussed “Chemical
inhibition of photosynthesis.” The speaker's abstract follows:

The rates of photosynthesis as measured by oxygen evolution in War-
burg manometers were determined with Chlorella vulgaris cells pretreated
with solutions of various inorganic compounds, and compared with control
rates. Several substances, including ZnSO4, CuSOs, (NH4)2SO4, H3BOs,
NiSOs, CoSOs, KCl, KI, and HgClz were found to inhibit photosynthesis,
whereas others like MnSO4, KNOs3, and MgSOz did not retard the process.
Inhibition was studied at five light intensities, from a range where light was
limiting to where it was in excess, in order to determine the effects of these
inhibitors on the photochemical and dark reactions in photosynthesis. A
comparison of control and pre-treated cell rates revealed differential inhibi-
tion. ZnSO, NiSOs, and KCl were found to inhibit the dark reaction with-
out appreciably affecting the light stage. CuSO4, H3BO3, and KI inhibited
the dark reaction but also retarded the light reaction to a lesser extent.
(NH)2SO4 and CoSOx4 caused a relatively equal inhibition of both reac-
tions. No substance was found which inhibited the light reaction alone.

The meeting adjourned at 4:40 P.M. to enjoy the delicious
refreshments provided by members of the Garden staff.
Respectfully submitted,

JOHN W. THOMPSON, JR.
RECORDING SECRETARY

MINUTES OF THE MEETING OF DECEMBER 2, 1941

The meeting was called to order by the President Dr. J. S.
Karling, at the American Museum of Natural History at 8:20 P.M.
Eighty-seven members and friends were present.

The minutes of the previous meeting were adopted as read.

Dr. Earl H. Newcomer, University of North Carolina, Chapel Hill,
N. C., was elected by unanimous ballot to annual membership.

The deaths of Professor W. J. Himmel, University of Nebraska, annual

member since 1924, and Mr. Severin Rapp, Sanford, Fla., associate member
since 1941, were announced with regret.

The President announced that the 75th Anniversary Committee
had selected the week of June 22, 1942, to hold the 75th Anniver-
sary Celebration meetings.

The scientific part of the program consisted of a talk by Pro-
fessor William Seifriz, of the University of Pennsylvania, on
“Recent advances in the study of protoplasm.” Professor Seifriz
illustrated his talk with motion pictures of the protoplasm of slime
molds.

The meeting was adjourned at 9:40 P.M.

Respectfully submitted,

JOLN Wey EOMESON ike
RECORDING SECRETARY

MINUTES OF THE MEETING OF DECEMBER 17, 1941

The meeting was called to order by the President, Dr. John S.
Karling, at 3:30 P. M. at the New York Botanical Garden. Fifty-
seven members and friends were present.

In the absence of the Recording Secretary, the Corresponding
Secretary read the minutes of the previous meeting. The minutes
were approved as read.

The following were elected by unanimous ballot to annual membership:
Mr. Russel Lee Walp, Marietta College, Marietta, Ohio; Miss Doris A.
Bach, 823 Park St., Kalamazoo, Mich.; Mr. Patrick Murray, St. Albert
College, Middletown, N. Y.; Miss Dorothy Day, Smith College, Northamp-
ton, Mass.; Miss Margaret S. Brown, 36 Kent St., Halifax, N. S.; Mr.
W. J. Nickerson, Harvard University, Cambridge, Mass.; Miss Clara S.
Hires, Mistaire Laboratories, 152 Glen Ave., Millburn, N. J.; Mr. Victor M.
Cutter, Cornell University, Ithaca, N. Y.; and Mr. D. G. Smith, 5 West
63rd St., New York, N. Y. To associate membership: Mr. I. E. Ehrenreich,

2944 West 28th St., Brooklyn, N. Y.; Rev. P. H. O’Neill, S.J., Fordham
University, New York, N. Y.; Miss Laura Filmyer, 2916 Grand Concourse,
New York, N. Y.; Miss Hope Mathewson, 82 East End Ave., New York,
N. Y.; Miss Margaret Fife, 82 East End Ave., New York, N. Y.; Mr.
Fred A. Buttrick, 184 Columbia Heights, Brooklyn, N. Y.; Miss Fairchild
Bowler, 1075 Park Ave., New York, N. Y.

The transfer of Dr. Hettie M. Chute, New Brunswick, N. J., from annual
to associate membership was approved.

The following resignations were accepted with regret: from annual
membership: Dr. Alfred S. Goodale, Amherst College; Miss Ernestine Ball,
Columbus, Ohio; Dr. Themistocles Acconci, Manhattan College; Mrs. D. C.
Boyce, Pittsburg, Pa.; Mr. Charles W. Slack, Atlanta Ga.; Dr. Arthur W.
Proetz, St. Louis, Mo.; Mr. G. M. Soxman, Dallas, Tex.; Miss Lena B.
Henderson, Lynchburg, Va.; Dr. J. E. Weaver, University of Nebraska;
Dr. J. W. Roberts, Beltsville, Md.; Dr. Valentine C. Baker, New York,
N. Y.; Miss Abigail O’Brien, Remsen, N. Y.; and Mrs. F. L. Keays, Great
Neck, N. Y.; from associate membership: Mrs. Cora Roe Smith, Branch-
ville, N. J.; Mrs. Regina Jais, New York, N. Y.; Mr. Spencer Scoit
Marsh, Madison, N. J.; Dr. Myrtle L. Massey, Brooklyn, N. Y.; Miss
Sarah J. Woodward, Brooklyn, N. Y.; Mr. Arthur E. Woods, East Orange,
N. J.; and Miss Ethelwyn Doolittle, New York, N. Y.

Dr. Robbins moved that Dr. Barnhart be delegated to repre-
sent the Torrey Botanical Club at the celebration of the 50th
Anniversary of the foundation of the Philadelphia Botanical Society
in Philadelphia on Friday, December 18, 1941. The motion was
seconded by Dr. Dodge and passed by the Club.

The scientific part of the program consisted of a talk and
demonstration on “Vitamins and growth of plants” by Dr. W. J.
Robbins of the New York Botanical Garden. The speaker’s
abstract follows :

It is now well established that the growth of many fungi is limited
by their inability to make adequate quantities of one or more vitamins.
Such fungi do not grow or grow poorly in a medium limited to pure sugars,
minerals and asparagine but on the addition of various substances of natural
origin or of one or more chemically pure vitamins, they develop satisfac-
torily. Ten species or strains of Ceratostomella were investigated.

The Cerastostomellas I used may be. grown readily in media to which
various natural products have been added, for example, malt agar, media
containing a decoction of tree bark, and so on. However, of the ten strains
or species reported here one only makes any considerable growth in a
medium limited to minerals, sugar and asparagine. This is Ceratostomella
pseudotsugae. However, the addition of vitamin B; and of vitamin Bg to
the medium materially increases the growth of that fungus. Biotin has no
effect. C. pseudotsugae shows a partial deficiency primarily for vitamin By

and secondarily for vitamin Bg. Ceratostomella piceaperda grows very slowly
in a medium of minerals, sugar and asparagine. The addition of biotin
and vitamin Bg markedly increases its growth. While C. pseudotsugae
evidences partial deficiencies for B,; and Be, C. piceaperda suffers from
partial deficiencies of biotin and Bg.

Ceratostomella ips isolated from Pinus ponderosa does not grow in the
basal medium. It suffers from a biotin deficiency and on the addition of
biotin to the medium grows quite satisfactorily. C. fimbricata and the
Ceratostomella from London Plane have a complete By deficiency. C. ulmi
has a nearly complete Bg deficiency. C. pini isolated from Pinus echinata
and C. pim isolated from Pinus ponderosa though differing somewhat in
appearance of growth are alike in having complete deficiency for both biotin
and By. C. montium and C. ips isolated from Pinus echinata suffer from
major deficiencies of By, Bg and biotin. They grow little or not at all unless
all three vitamins are present in the medium. Among these ten species or
strains of Ceratostomella seven different types of vitamin deficiencies exist:

1. Major or complete deficiency for By—little affected by Bg or biotin.
Major or complete deficiency for Bg—little affected by By or biotin.
Major or complete deficiency for biotin—little affected by Bi or Be.
Partial deficiency B; and Bg—little affected by biotin.

Partial deficiency biotin and Bg—little affected by By.

Major deficiency biotin and B,—little affected by Be.

Major deficiency B;, Bg and biotin.

ow ON Ga gS GO)

By selecting a suitable species of Ceratostomella it is possible by its
growth or failure to grow to demonstrate the presence or absence of Bi, Be
or biotin or substitutes therefor. In the course of these experiments it was
discovered by accident that an extract of cotton batting added to a medium
of minerals, sugar and asparagine permitted good growth of Ceratostomellas
which showed deficiencies for By, Bg or biotin or combinations. It seems
justifiable to conclude that unbleached and unwashed cotton contains sig-
nificant quantities of all three of these vitamins.

In the same way, that is by the growth of various species of Ceratosto-
mella, the presence of By, Bg and biotin in unpurified Difco agar also was
determined.

Since cotton and Difco agar are both commonly used in laboratory
procedures, it is clear that due consideration must be given to them as pos-
sible sources of vitamins. Knight and his associates working with the
so-called sporogenes vitamin found that stray filaments of cotton falling in
their media invalidated their bacterial experiments.

In presenting these results I have emphasized the more marked deficien-
cies of the Ceratostomellas for the three vitamins By, Bg and biotin. Less
marked deficiencies have been observed. For example, a species which grows
little or not at all unless By is added to the medium may grow somewhat
more rapidly if all three vitamins are added. It is probable also that some
‘of these organisms suffer from partial deficiencies for other vitamins or
vitamin-like growth substances. I am not sure that reproduction will occur

in media supplemented with B1, Bg and biotin as satisfactorily as it does in
media containing natural products, for example, malt agar. Some evidence
for the deficiencies for unidentified growth substances is furnished by the
more rapid growth in some natural media than in a basal medium containing
twelve pure vitamins and twenty-one pure amino acids.

These results with Ceratostomella are of interest:

1. Because of the diversity of vitamin deficiencies in representatives
of a single genus.

2. Because the discovery of a fungus with nearly complete vitamin Bg
deficiency suggests that it may be used for bio-assay for this
vitamin. Assay methods for vitamin Bg are at present unsatis-
factory.

3. Because of the determination of the presence of significant quan-
tities of biotin, Bg and By, in cotton batting and unpurified
agar.

4. Because the results show that a fungus may suffer from a com-
plete deficiency of three vitamins, a situation which approaches
more nearly the condition of the animal where many complete
deficiencies exist. This emphasizes again the fundamental like-
ness of the basic physiological processes in all living things.

The meeting was adjourned at 4:20 P.M. and many members
and guests remained to continue the discussion informally at tea
provided by the Garden staff.

Respectfully submitted,

JOHN W. THOMPSON, JR.
RECORDING SECRETARY

THe FreLp CoMMitTeEE of the Club announced 168 botanical events
in its schedules during 1941. Of these 85 were actual field trips,
many of them in cooperation with one or more other botanical
societies. Reports were received from 78 of these field trips. Total
attendance was 1456 or an average of about 19 persons to each
field trip. The high mark was the Branchville Nature Conference,
attended by 97.

THE TORREY BOTANICAL CLUB
OFFICERS FOR 1942

President: C. STUART GAGER
Vice-Presidents: JoHN A. SMaAtt, F. Treasurer: WW. GorDON WHALEY
CLYDE CHANDLER

; Editor: Harotp W. RIcKETT
Recording Secretary: Miss Honor M.

HoLLINGHURST Business Manager: MicHAEL LEVINE
Corresponding Secretary: Harotp C. Bibliographer: Mrs. LAzeELtta ScHWAR-
Bop TEN

Delegate to the Council, N. Y. Academy of Sciences: W. J. Rospins

Representatives on the Council of the American Association for the
Advancement of Science:

A. E. HircHcock J. S. Karine
Representative on the Board of Managers of the N. Y. Botanical Garden:
H. A. GLEASON

Council for 1940

Ex officio members

Bernard O. Dodge Edwin B. Matzke Michael Levine
Arthur H. Graves i Harold N. Moldenke William J. Robbins
Alfred Gundersen John S. Karling Henry K. Svenson
George T. Hastings Florence C. Chandler John A. Small

Harold W. Rickett

Elected members

1939-1941 1940-1942 1941-1943
Gladys P. Anderson Lela V. Barton Helen M. Trelease
John M. Arthur Ralph H. Cheney Ralph C. Benedict
Harold H. Clum Robert A. Harper John H. Barnhart
Percy W. Zimmerman Edmund W. Sinnott

Committees for 1940
ENDOWMENT COMMITTEE

Clarence Lewis, Chairman J. Ashton Allis
Caroline C. Haynes Henry de la Montagne Helen M. Trelease
ProGRAM COMMITTEE
Harold C. Bold, Chairman (e-officio) E. Marcy
William J. Robbins P. W. Zimmerman
E. B. Matzke G. Whaley

FIELD COMMITTEE
John A. Small, Chairman

Edward J. Alexander Rutherford Platt Robert Hagelstein
G. G. Nearing Henry K. Svenson Michael Levine
Vernon L. Frazee Ellys Butler James Murphy
Alfred Gundersen Dolores Fay Daniel Smiley, Jr.
Inez M. Haring Eleanor Friend Farida A. Wiley

H. N. Moldenke

LocaLt Frora COMMITTEE
W. H. Camp. Chairman

William J. Bonisteel Harold W. Rickett Dolores Fay

James Edwards Ora B. Smith H. Allan Gleason

John M. Fogg, Jr. Herbert M. Denslow Hester M. Rusk
Cryptogams

Ferns and Fern Allies: R. C. Benedict, W. Herbert Dole, N. E. Pfeiffer
Mosses: E. B. Bartram

Liverworts: A. W. Evans, E. B. Matzke

Freshwater Algae: H.C, Bold

Marine Algae: J. J. Copeland

Fungi: A. H. Graves, J. S. Karling, W. S. Thomas

Lichens: J. W. Thomson, Jr.

Myzomycetes: R. Hagelstein

CoMMITTEE ON EXCHANGES
Harold C. Bold Amy Hepburn Elizabeth Hall

OTHER PUBLICATIONS

OF THE

TORREY BOTANICAL CLUB

(1) BULLETIN

A journal devoted to general botany, established in 1870 and
published monthly, except during July, August, and September.
Vol. 68, published in 1941, contained 694 pages of text and 55 full
page plates. Price $6.00 per annum. For Europe, $6.25.

In addition to papers giving the results of research, each issue
contains the INDEX TO AMERICAN BOTANICAL LITERATURE—a very
comprehensive bibliography of current publications in American
botany. Many workers find this an extremely valuable feature of the
BULLETIN.

Of former volumes, 24-68 can be supplied separately at $6.00
each; certain numbers of other volumes are available, but the entire
stock of some numbers has been reserved for the completion of sets.
Single copies (75 cents) will be furnished only when not breaking
complete volumes.

(2) MEMOIRS

The Memoirs, established 1889, are published at irregular in-
tervals. Volumes 1-18 are now completed. Volume 17, containing
Proceedings of the Semi-Centennial Anniversary of the Club, 490
pages, was issued in 1918, price $5.00.

Volume 18, no. 1, 108 pages, 1931, price $2.00. Volume 18, no.
2, 220 pages, 1932, price $4.00. Volume 18 complete, price $5.00.

Volume 19, no. 1, 92 pages, 1937, price $1.50. Volume 19, no.
2, 178 pages, 1938, price $2.00.

(3) INDEX TO AMERICAN BOTANICAL
LITERATURE

Reprinted monthly on cards, and furnished to subscribers at three |
cents a card.
‘Correspondence relating to the above publications should be
addressed to
W. Gordon WHALEY,
Barnard College,
Columbia University,
New York, N. Y.

Heroariun, LIBRAR

Volume 42 March-April, 1942 Number 2"FW YO

COTANI¢
GARDE
A Bi-MonTHLy JouURNAL oF BotanicaL Notes anp News
EDITED FOR
THE TORREY BOTANICAL CLUB
BY
WILLIAM J. BONISTEEL
John Torrey, 1796-1873
CONTENTS

A Botanist’s Summer in Costa Rica........................ M. A. CurysLerR 33
Collecting Chicle in the American Tropics (Part 1)......... Joun S. Kariine 38
Rare Cladonia in New Jersey............. 0000s ccc ee cence cca eeeas W. L. Dix 49
Book Reviews

Introducing (Insects iy. i5 2 kel d acd pie) aus ede k Slated oa) cerned JAMES Forses 50

The Flower Family Album.......................... GrorceE T. Hastines 51

ISU LESTE TES 555 is Ha we ent eh es OE Oey ae LC CE GeEorcE T. Hastincs 52

Bloray of indiana) 75 Os es ae Gees OE oa diate aia hele, ate R. M. Harper 53
Field) Drips of the Glubiar ara coun fo dd Seater eA SOM Mes uses at cnt! 56
Proceedings’ of the (Clube (oe eis peta ale HN Oe SRN AA 61
News sNOteSe tte et Rare Mew UTES) Des NEI DAM a Ue NaS ea Ul MSL iS aa Mave te 64

PUBLISHED FOR THE CLUB

By THE FREE Press PRINTING COMPANY
187 CoLLEGE STREET, BURLINGTON, VERMONT

Entered as second class matter at the post office at Burlington, Vermont,
October 14, 1939, under the Act of March 3, 1879

TORREYA

TorrEYA, the bi-monthly publication of the Torrey Botanical Club, was established
in 1901. Torreya was established as a means of publishing shorter papers and inter-
esting notes on the local flora range of the club. The proceedings of the club, book
reviews, field trips and news notes are published from time to time. The pages of
TOoRREYA are open to members of the club and others who may have short articles
for publication.

TorREYA is furnished to subscribers in the United States and Canada for one
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elsewhere, twenty-five cents extra, or the equivalent thereof. Postal or express
money orders, drafts, and personal checks are accepted in payment. Subscriptions are
received only for full volumes.

Of the annual membership dues of the Torrey Botanical Club, $.50 is for a year’s
subscription to TORREYA.

INSTRUCTIONS TO CONTRIBUTORS

Drawings and photographs should be mounted on stiff cardboard and the desired
reductions plainly indicated. Figures should be so planned that after reduction they
will occupy the entire width of a page (4 inches) and any portion of the height
(6%4 inches). Labels should be parallel to the shorter dimension of the page. It is
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cluded with the manuscript (not affixed to the figures). All legends for one group
of figures should form a single paragraph. If magnifications are stated, they should
apply to the reduced figures.

TorreYA is edited for the Torrey Botanical Club by

WM. J. BONISTEEL
W. H. CAMP DOROTHY J. LONGACRE

MEMBERSHIP IN THE TORREY BOTANICAL CLUB

All persons interested in botany are invited to join the club. There are four
classes of membership: Sustaining, at $15.00 a year; Life, at $100; Annual, at $5.00
a year and Associate, at $2.00 a year. The privileges of members, except Associate,
are: (a) To attend all meetings of the club and to take part in the business, and -
(b) to receive its publications. Associate members have the privilege of attending
meetings, field trips and of receiving the Schedule of the Field Trips and the Bulletin
of the New York Academy of Sciences.

Manuscripts for publication, books and papers for review, reports of field
trips and miscellaneous news items should be addressed to:

DR. WM. J. BONISTEEL
BIOLOGICAL LABORATORIES, FORDHAM UNIVERSITY
New York, N. Y.

TORREYA

Wor 42 MarcuH-APRIL No. 2

A Botanist’s Summer in Costa Rica

M. A. CHRYSLER

It was the writer’s good fortune to spend July and August of
1940 in the little republic of Costa Rica, which has been character-
ized by Gunther’ as “‘one of the most delightful countries in the world
and one of the purest democracies on earth.” It has moreover a par-
ticularly interesting flora, especially to the student of ferns. Accord-
ing to the North American Flora, it is headquarters for Gleichenta-
ceae, with an array of endemic species, hence a trip was arranged so
as to provide a two-weeks’ stay in Jamaica, a week on Barro Colo-
rado Island, C. Z., and the balance of the season in Costa Rica. Dur-
ing most of the time the writer was accompanied by his colleague,
Dr. W. E. Roever, whose cooperation was invaluable.

One gains a lasting impression of the vertical distribution of the
plant life by looking out of the window from the train which takes
him from Port Limon to the capital, San José—a trip of only a
hundred miles which nevertheless occupies about six hours. Starting
from the banana groves near the coast, the traveler passes through
real jungle with reappearance of bananas, coconuts and cacao at
every settlement—the tierra caliente. Presently the lower stretches
of the Reventazon River are reached, and the road begins a series of
sharp curves and steep grades as it follows the course of the rushing
river. By the time an elevation of 2000 feet is reached, coffee has re-
placed the banana as the leading crop, giving an entirely different
aspect to the landscape, for the coffee shrubs grow in the partial
shade of such trees as species of Inga, and during August are bright
by reason of the ripe red berries which contain the familiar coffee
“bean.” The railroad banks are enlivened by the brilliant flowers of
Heliconia and Costus, representing monocotyledonous families quite
unknown to northern floras.

1 Gunther, John. Inside Latin America. Harper & Brothers, New York,
1941.

Torreya for March-April (Vol. 42: 33-64) was issued April 10, 1942.
$8}

We are now passing into the tierra templada of Standley,” the
region in which most of the population is found. The curves become
sharper and the grades if possible more steep, as we realize when a
brisk shower descends and the track becomes so slippery that the
train is stalled until the rails are sanded and the plucky little engine
jerks the train into motion, while we breathe more easily although
we realize that perhaps we should have bought some of that fried
chicken which was offered at the car windows while we stopped at
Turrialba. The view of river and mountains grows more expansive,
and Roever’s Leica is in frequent use. At length an altitude of 5137
feet is attained at the Continental Divide just beyond Cartago, the
former capital, which was levelled by an earthquake thirty-odd years
ago. The train slides down the remaining ten miles to San José,
situated at an altitude of 3800 feet among the coffee plantations in
the saucer-shaped “meseta central.”

San José was our headquarters for most of the two months, and
was convenient because of the bus lines radiating in every direction.
Under the guidance of Director Valerio and Dr. A. Skutch of the
Museo Nacional, we made our first excursion to the tierra fria, go-
ing by auto on one of the few paved roads until an elevation of 6800
feet was reached, where we found the way blocked by a landslide.
So we finished on foot the few miles to the hamlet called Varra
Blanca, where we spent a memorable week. Here no crops except
potatoes are raised, and the universal industry is dairying. Milk,
tortillas, beans and rice are the staple articles of diet. As we wand-
ered out into the fields we were at once attracted by huge pink
bouquets formed by old oak trunks covered with climbing shrubs
belonging to the ericaceous genus Cavendishia. The dominance of
epiphytes astonished us until it was realized that these plants en-
joyed plenty of light, air and water, also immunity from grazing
animals. Every tree had its assortment of “air-plants,” chiefly ferns
and orchids. One large shaggy species of Trichomanes (T. lucens)
attracted attention, also what appeared to be a fleshy spleenwort
(Enterosora spongiosa). One tree was beautifully mantled by a
vigorous specimen of the familiar Polypodium aureum, below which
a border of Nephrolepis pendula was added by way of good measure.
The fragrant Asplenium auritum adorned the base of most trees,

2 Standley, P. C. Flora of Costa Rica, part 1. Chicago, 1937.

while the dainty Rhipidopteris peltata grew in masses on fallen
trunks. Presently the usual afternoon shower drove us to cover,
where we hastened to get our treasures into the drier or into pickle
ere the quick tropical night descended and we had to depend on
candle light.

The scientific peak of the whole trip was reached when on a hill-
side near our stopping place we found eight species of Dicranopteris
(a segregate of Gleichenia) including the endemic D. costaricensis
and the remarkable D. retroflexa. D. Bancroftu (Fig. 1) afforded
a surprise, for instead of the single fork bearing two leaflets found

Ficure 1. Dicranopteris Bancroftii filling a small ravine; the branches of the
leaves are two feet long. Varra Blanca, C. R. Alt. 6000 ft.

earlier in Jamaican plants, forks of the second, third and even fourth
order were characteristic of the plants in a ravine near those endemic
species. Stream banks displayed a huge herbaceous Senecio (Coo-
peri) and an equally large Eupatorium (angulare) while the
fuchsia used as a house plant was represented by F. arborescens 15
feet high. Melastomes of various genera—trees, shrubs, and herbs
—were of constant occurrence, some as beautiful as climbing roses
(Blakea spp.). From Varra Blanca Dr. Roever took a memorable

trip to the crater of Volcan Poas (8300 feet), bringing back Drimys
Wintert, famous because although it is a dicotyledon its wood shows
tracheids in place of vessels. Other prizes were certain rare ericads
and the immigrant conifer from South America, Podocarpus mon-
fanus.

Our next trip afforded a chance to sample the rich flora of a
region at an altitude of 2,200 feet, San Isidro del General. This vil-
lage in the midst of a bean growing region has the distinction of
having skipped some of the usual evolutionary stages of a com-
munity, such as horse and carriage, automobile, railroad, telegraph
and telephone, for it has leaped from the ox-cart stage to airplane
and radio. Half an hour by plane covered the journey from San
José, although by mule-back over the ridges five days used to be
consumed. We were particularly impressed at San Isidro by the
variety of tree ferns and the pendent species of Lycopodium. Al-
though the roadsides showed some highly colored flowers, the only
conspicuous angiosperms in the rain forest were the orchid-like
Orchillium Endresii—a large-flowered member of the bladderwort
family—and Cephaelis spp. (Rubiaceae) distinguished by two deep
red bracts enclosing each inflorescence. But the Selaginellas of
stream-banks, a splendid Lindsaya (lancea) a climbing Blechnum,
impressive Dennstaedtias made up for paucity of color.

Another area along the 2,000-foot contour was visited—the val-
ley of the Reventazon River near Turrialba village. The calcareous
banks of the river support a varied flora, again consisting chiefly of
ferns, but including Zamia Skinnert, a species interesting because of
its trunk-forming habit. We were hospitably entertained at a coffee
plantation by Mrs. Goode, the patron saint of botanists in that re-
gion, where every hedge-row presents novel plants, and a bewilder-
ing assortment of Dryopteris challenges one’s observing capacity.

The vicinity of Cartago has been made familiar to biologists by
Professor and Mrs. Calvert* through the notable volume describing
a year’s work, chiefly on insects but containing many references to
plants. Although the region is much changed during the last thirty
years, we still found the slopes of Mt. Carpintera well worth explor-
ing, while the thickets and walls could be depended on to furnish

= Calvert, P. P. and A. C. A year of Costa Rican natural history. New
York, 1917.

unfamiliar ferns. The chief attraction of the region, however, is the
orchid garden of Mr. C. H. Lankester. This is a most remarkable
assemblage of orchids of Costa Rica and other tropical countries,
blended with ferns and cycads and growing on trees and banks in a
charming atmosphere of wildness. The writer was most kindly en-
tertained by the Lankesters, and cannot forget the display of hybrid
orchids, some of rich fragrance, which greeted him each morning as
he emerged from sleeping quarters. The visit was notable for a
number of personally conducted field trips, one to a station for
Ophioglossum palmatum and another along the newly constructed —
Pan-American highway.

The difficulties experienced in travelling in Costa Rica are il-
lustrated by another trip. In company with Sr. Leon of the museum
we took bus for Heredia, then climbed six miles on a dirt road to
the slopes of Volcan Barba. On the way a sudden storm overtook
us, as we ate lunch by a friendly bank. Arriving at a schoolhouse we
went under the guidance of the schoolmaster to a sulphur spring
near which we collected a Peperomia which is regarded as a new
species. We were allowed to sleep on the floor of the schoolhouse,
which we may report as clean and polished, but cold and drafty. It
was on this trip that we found Botrychium cicutarium, B. under-
woodianum and Ophioglossum reticulatum, at altitude 6,500 feet.
It was in a similar locality that we found Gunnera insignis, a plant
provided with leaves so large that they are used as umbrellas by the
natives.

Our advisers did not encourage us to brave the dangers of ma-
laria by venturing into the Province of Guanacaste, on the Pacific
slope. The climate of San José is so healthful, and relatively easy
excursions are so many that our remaining trips were made in the
neighborhood of the capital. On the last day of August we took train
for Port Limon, feeling content at having accomplished at least the
main objects of the trip.

DEPARTMENT OF BOTANY
RutTGerRsS UNIVERSITY

Collecting Chicle in the American Tropics*

Joun S. KARLING

The principal source of chicle, the basic ingredient of chewing
gum, is the latex of Achras zapota, a species of the family Sapotaceae
which occurs in abundance in southern Mexico and Central Amer-
ica. The sapodilla or chicle tree is generally regarded as indigenous
to southern Mexico, Central America, northern South America, and
the West Indies, but because of its delicious fruit it has been planted
extensively and may now be found under cultivation in limited quan-
tities as a fruit tree in most tropical and sub-tropical countries. It is
principally in southern Mexico and Central America, however, that
it grows in sufficient quantity, size, and height to make tapping for
chicle profitable. Here the trees may occasionally attain a height of
a hundred feet with straight smooth boles, sometimes as much as
eight to twelve feet in circumference; and in these regions during
the past half century has sprung up the extensive and unique indus-
try of gathering crude chicle which has no parallel in any other part
of the world.

Although the natives in tropical America had been using small
amounts of chicle for various purposes in pre-Columbian times
(Melendez, 1920), it was not until the discovery of chicle as a suit-
able base for chewing gum that this product became economically
important. This discovery more than half a century ago is said to
have been the result of attempts to vulcanize the gum of the sapodilla
tree in the same manner and as a possible substitute for rubber. The
similarity of chicle to spruce and cherry gums, the best chewing
gums in use at that time, and its adaptability to chewing and com-
pounding with adulterants, sugars, and flavors were soon recognized,
and from these first modest experiments and an initial outlay of fifty-
five dollars the extensive present-day chewing gum industry is said
to have had its beginning. Hand in hand with the spread of the gum
chewing habit grew the demand for raw chicle, and within a few
years a new enterprise sprang up in the jungles of southern Mexico
and Central America. Rival American contractors began to push into
the jungles to obtain large concession of virgin forests and to offer
unheard-of inducements to the natives for gathering chicle. Raw

1 Address presented before the Torrey Botanical Club, December, 1940.

3!)

chicle thus soon became one of the principal exports of several Mexi-
can and Central American states, and in 1930 the import of chicle
into the United States had risen to nearly fourteen million pounds
(U. S. 1932). In its half century of growth the chewing gum in-
dustry has made phenomenal progress, and at the present time ranks
among the big American industries. The manufactured output in
1930 was valued at more than seventy million dollars, representing
a retail business of over a hundred million dollars.

CHICLEROS AND THE PRESENT NATIVE METHOD OF
TAPPING AND PREPARING RAw CHICLE

The native laborers or Indians who bleed the sapodilla trees and
gather the chicle are known as chicleros. No particular group or
tribe of natives has a monopoly of skill in this profession, and chic-
leros of almost every race, color, nationality, and intermixture are to
be found. The native Indian of southern Mexico and Central Amer-
ica, however, is generally regarded as the most skillful, careful, and
desirable. Steadiness of hand and accuracy in manipulating a ma-
chete as well as a certain amount of skill in climbing are the prime
requisites of a good chiclero, and only a small proportion of the na-
tive labor is capable of bleeding chicle. The chiclero is thus regarded
as a skilled worker in the tropical forests and is among the best paid
of all native laborers. Since the chicle tapping season is dependent
on the rainfall, the chicleros spend the greater part of the rainy
season from July to February in the chicle forests. As soon as the
tropical rains start in June, the exodus of chicleros from the coast
towns and villages into the jungle of Peten, Quintana Roo, Yucatan,
Campeche, Chiapas, and British Honduras begins. For the purpose
of companionship as well as assistance in certain aspects of the work,
they generally go in small groups of from two to five, and may often-
times take their families with them to form large camps.

While in the chicle forests the chicleros live in temporary camps
close to the scene of operation. These camps are generally located
on the edge of a lagoon, swamp, or savannah where water is avail-
able, since a constant water supply is necessary not only for drinking
and cooking but for the molding of cooked chicle as well. As a con-
sequence, the camps are generally situated where mosquitoes and
other blood-sucking insects are likely to be most numerous. In fig-
ures 4 and 5 is shown a portion of such a chicle camp at the border

M. Hedler.

. Photograph by H.

s zapota for chicle

Achra

ing

Chicleros tapp

of a dense cohune palm ridge. Inasmuch as the chicleros may often
shift their operations during the same season and rarely return to
the same camp in successive years, their huts are but temporary
structures of upright poles in the ground roofed over with palm
leaves. During prolonged tropical rainstorms these huts afford but
little protection from the rain, so that the chicleros are more or less
wet for extended periods of time. This constant exposure, together
with the presence of disease-carrying insects, often leads to the con-
traction of malaria and other pernicious tropical diseases. While in
the jungles the chiclero’s fare is very simple and consists chiefly of
rice, frijoles, and tortillas, with occasional meat from wild game
which they may kill.

The chicleros are paid in accordance to the amount of gum ex-
tracted during the chicle season. A skilled tapper in a virgin forest
can sometimes collect as much as 2000 pounds in one season, for
which he is paid from 12 to 30 cents per pound, depending on the
quality and moisture content of the gum. Since he is frequently able
to make more money gathering chicle during the rainy season than
for working for wages throughout the entire year, he is preferably
idle from February to June. During this period many of them loaf
from one village to another doing an occasional job. As a result they
may be partially or wholly dependent on some chicle contractor for
rations and livelihood during the dry months, and by the time the
chicle season arrives they are often in debt for more than the value
of the chicle they can extract. In this manner, they often become
bound to one contractor from year to year. For these reasons many
observers maintain that the chicle industry has done more harm than
good to native labor in tropical America. It has been claimed that
previous to the advent of chicle the Indian was a fairly industrious
and conscientious worker who cultivated his milpa, hunted, and was
quite contented to work for small wages rather than remain idle,
while the women spun and made their own clothing. Then came the
American chicle contractor offering fabulous advances in cash for
chicle with the inevitable result that the Indian forsook his milpa to
become a chiclero. Finding that he was thus able to more than double
his yearly income in a few months gathering chicle, he refused to
work at all during the dry season or only for greatly increased wages.
As a result the milpa was neglected, and the price of rice, beans, and
corn nearly doubled. Moreover, the earning of more money than

‘purying ‘qd “MM 4q syderdojoyg
‘g]o1Yy9 Joy vjodez sesyoy surdde, Jo poyzou Jetids-ayoyoeul dATyeU oY,

nrpemrmer ements tame eT ATTN

was necessary for food during the chicle season developed extrava-
gant tastes among the women. Where once they had been contented
with the simple native costume, they now demanded expensive cloth-
ing, etc., from their men. ‘The increased price of food and the efforts
to satisfy more extravagant tastes were not commensurate with
their increased earnings from bleeding chicle, and as a result they
had to turn continually to some chicle contractor for advances. He,
in turn, charged impossible rates of interest and paid as little as pos-
sible for the chicle. The chiclero, not to be outdone, adulterated the
chicle, and often received advances from several contractors without
working for any of them. In the last two decades, however, many of
these economic factors have changed considerably.

The chicle tapping season is dependent on the rainfall, and is
thus concurrent with the rainy season, contrary to the reports of
certain authors. If the tropical rains come early it may begin in June
and extend to February, but it does not generally get well started
until July and August. Tapping commences with the daylight. The
chiclero rises while it is yet dark, prepares a light breakfast, and
starts out afoot through the dense jungle for the sapodilla trees
which he had located on previous days. Arrived at a tree, the chiclero
first clears a small area around the tree, and adjusts the skin or can-
vas bag in which the milk is collected. It is either set on the ground
or hung from an incision in the bark (Figs. 1, 2). Directly above the
bag a small area of the tree is cleared of its outer hard bark and an
upward incision made in the softer cortex with the machete. This
makes a flap under which the end of a trimmed palm leaf is inserted
to act as a conveyor for the latex from the tree to the bag. Having
properly adjusted the bag and inserted the palm leaf, the chiclero
begins to tap.

Tapping in the wild chicle “bush” or “chicleria” is done exclu-
sively with a long thin-blade cutlass or machete. Chicleros generally
prefer the fairly straight machete, since they are thereby able to re-
move a wider chip of bark with each stroke. The native method of
tapping is essentially a half-spiral system, as is shown in Figures
2 and 3, and consists of successive parallel rows of cuts ascending
the bole obliquely. The successive oblique rows of incisions alternate
from side to side and lead into the lower preceding ones, so that the
latex from the individual rows flows together in a zigzag channel
down the tree to the point where the collecting bag is attached. As

ish Honduras.

in Bri

a
=
3
O
n

iclero

A ch

is at once apparent in Figures 2 and 3, the obliquely ascending rows
of cuts are not continuous channels encircling the trunk, but consist
or more or less separate partially overlapping incisions. The latex
from each cut thus spills over into the lower preceding one. The first
oblique rows are usually started a few inches above the point of in-
sertion of the palm-leaf conveyor, at angles of 45° to 70°, depending
on the size of the tree and the habits of the chiclero, and the follow-
ing oblique rows are then made ten to twenty inches apart. In the
case of large and old trees, the outer bark is generally too hard and
thick for making satisfactory incisions, and as a consequence it is
usually removed before each oblique channel is made. This is well
shown by the tree in the foreground of Figure 1 and in Figure 2. In
the small tree shown in Figure 3, however, removal of the outer
bark was not necessary. Chicleros thus frequently carry two ma-
chetes, an old one for removing the outer bark and another sharper
one for making the oblique incisions.

As soon as the chiclero has tapped as high as he can reach stand-
ing on the ground he begins to climb. This is done with the aid of
a thick rope passed around the tree and the middle of his body. The
looseness of the loop permits the chiclero to steady himself with his
feet against the tree, leaving both hands free for tapping, as is illus-
trated in Figure 1. The trees shown in this figure are being tapped
for the second time, and the overlapping of the previous and the new
oblique channels accounts for the striking diamond-shaped areas on
the bole. Climbing spurs such as those used by telephone linemen
are frequently employed as an aid, but the best chicleros spurn such
assistance and climb only with bare feet. The chiclero thus climbs
higher and higher, swinging from side to side as he makes the alter-
nate rows of cuts until the entire bole has been tapped. In cases
where the trunk forks and large erect branches are present, the
latter also may be tapped. When one tree is finished the chiclero
proceeds to the next, and so on into the early forenoon until relative
humidity, sun, wind, and temperature begin to affect the flow of
latex. In the dense jungle where only slight winds penetrate relative
humidity remains fairly high, so that tapping may continue until
middle forenoon. In the more open “bush,” however, increased tem-
perature, sun and wind, and loss in humidity make tapping unprofit-
able after 8 or 9 o’clock in the morning.

The chiclero usually spends the remainder of the forenoon locat-
ing trees for the next day’s tapping. The first step after finding a
virgin tree is to test its flow of latex. Incisions are made in the bark
near the base, and if the latex flow is good the tree is marked or
staked for tapping; otherwise it is discarded. In an area which has

Aerating chicle after cooking.

once been tapped, it is not uncommon to find several large, vigorous,
and sound trees that are untouched. They all, however, bear the test
marks of the chiclero, testimony to the fact that they are poor yield-
ers. By the careful selection thus of only good yielding trees, the
amount of chicle per tree in the virgin sapodilla forests may be quite
high, but the acreage yield is proportionately low. The yield per in-
dividual tree is quite variable, as is to be expected in a wild popula-
tion. Some trees do not yield sufficient latex to wet the incisions,
while others have been reported to yield as much as sixty-one pounds
(Hummel, 1925). The report of Sperber that trees in Mexico yield
thirty to thirty-five pounds annually is obviously without foundation.
Exceptionally large trees may yield that amount at the initial tap-
ping, but certainly not every year. According to the writer’s obser-
vations, the initial yield per large virgin tree is usually two to ten
pounds. As soon as a sufficient number of trees have been located,
the chiclero returns to camp with the bags of latex. Frequently the
bags are allowed to remain on the trees until the following day if
there is no danger of rain. Otherwise they are collected on the same
day, since the presence of excess water in the latex makes cooking
long, tedious, and difficult.

The latex from the various trees is poured together in empty
petrol tins or larger bags and stored until a sufficient amount for
cooking has been accumulated. The chicleros generally tap through-
out the week and cook the latex on Sundays. Cooking is primarily
for the purpose of driving off water and is done in large iron kettles
or cauldrons over an open fire, as is illustrated in Figures 4 and 5.
The time required for cooking varies with the amount and quality
of latex in the pot, but usually one to two hours are sufficient. Dur-
ing the process the latex is stirred continuously with a long stick or
paddle in a circular fashion to prevent burning and to throw the
water toward the periphery of the mass. After the latex has reached
the consistency of soft taffy, the fire is scraped from beneath the
kettle or the latter is removed from the open fire. The gum is then
further worked and aerated (Fig. 6) with the long paddle until it
begins to cool and become firm. It is then lifted out onto a large
soaped palm leaf, tarpaulin sheet or sack (Fig. 7) and molded into
blocks (Fig. 8). The chicle at this stage is still quite sticky, and an
abundance of water and soap are essential for successful handling
and molding. Preparatory to taking the chicle out of the cauldron,

the chicleros soap their hands and arms thoroughly, and by constant
renewal of soap they are able to handle and mold it with a minimum
of sticking. In the earlier days of gum collecting the chicle was often
heaped together into semi-spherical and rectangular masses and al-
lowed to dry, but the general practice at present is to mold it in

Molding chicle into blocks.

rectangular frames of uniform size, which allows greater economy
of space in storing and shipping. Formerly, the weight of the chicle
masses and blocks varied as much as their shape, and blocks weigh-
ing as much as a hundred pounds were not uncommon. At present,
however, they are usually more uniform in weight, varying from
twenty-five to forty pounds. At the time of molding the chicle con-
tains approximately forty to fifty per cent water, but by the time it
is shipped to the States, the moisture content varies from twenty-
five to thirty-five per cent.

As soon as the chicleros have accumulated a fair number of
blocks, the chicle is delivered to the contractors who advanced the
money for rations and supplies. In more organized chicle operations
the contractors generally send out pack mules periodically to the
various chiclero camps to collect the accumulated chicle. It is then
concentrated in central camps, baled and transported by mule, boat
or aeroplane to storehouses along the coast, and eventually shipped
to the United States or Canada.

Attempts have also been made from time to time to extract chicle
profitably from the leaves and green fruit, but without much suc-
cess. It is estimated (Anonymous, 1923) that approximately 3200
leaves are required for a pound of gum, and the cost of production
at that rate is in excess of the present price of chicle.

(To be continued)

Rare Cladoniae in New Jersey
W. L. Dix

Cladonia squamosa {. carneopallida Sandstede. This specimen
was collected by the writer a few miles east of Jackson, Camden
County. It was first collected in America near Hartford, Connecti-
cut, and reported by Dr. Alexander W. Evans in his third supple-
ment to the Cladoniae of Connecticut.

Cladoma pyxidata var. neglecta f. centralis Schaer. In this form
the cup is centrally proliferate. It was collected near Hopewell in
Mercer County. Apparently this is the first account of this form in
America.

The second authenticated collection of Cladoma turgida in New
Jersey was made by the writer along the Appalachian Trail in War-

ren County last November. The reported collections of Austin from
Bergen County, by Eckfelt from Warren County, and by Torrey
from Passaic County, all proved to be other species, or are repre-
sented by no specimen. Torrey, however, did later find “a small form
of the species” in Green Bank State Forest, Atlantic County, in
1936. The specimen collected by the writer consisted of squamules
only.

BOOK REVIEWS
The A. B. C. of Insects

Introducing Insects: A Book for Beginners. By James G. Needham. The
Jaques Cattell Press, Lancaster, Penna. 1940. Pp. v + 129. $1.50.

Biting into a wormy apple is an unpleasant experience ! Unpack-
ing the dress suit after it has been stored for a while only to find
that it has a few conspicuous moth holes can be very distressing. If
we know nothing about these “pesky bugs” that come to upset us or
if we should like to refresh our memories, Professor Needham’s little
book is an ideal starting point.

Professor Needham has given us a book, written in simple, non-
technical language coupled with an easy style, that will help the lay-
man to understand and to appreciate the insects with which he comes
in contact. His introductory chapters, “Why Study Insects” and
“How to Study Insects,” are followed by discussions of such com-
mon insects as butterflies, dragonflies, grasshoppers, leaf bugs,
beetles, scale insects, aphids, mosquitoes, and bees. The author next
considers such ecological groups as carniverous insects and insects
that eat our foods and textiles. The concluding chapters are con-
cerned with the control of insects and the collecting and rearing of
insects.

The expositions of life histories, food habits, and habitats pre-
ferred by the insects during their developmental stages are inter-
spersed with considerations of the balance in nature, parasitic and
predatory insects, beneficial and noxious insects. The importance of
careful observation is stressed so that we shall know which insect to
swat and which insect to protect: our insect friends should not be
needlessly killed.

The line drawing illustrations are well chosen and admirably
compliment the various parts of the text. In illustrations of this type,

most drawings have to be larger than the actual specimen under con-
sideration to show even a minimum of structure. As the book is
especially designed for beginners, it is to be regretted that hairlines,
indicating the size of the insects, are not included with all the draw-
ings.

This book can be used with profit in nature study groups or clubs
and in biology classes which range in age from the early teens
through adults. The suggested points for field observations, tricks
for catching some of the insects, the study of live insects, and the
simple rearing experiments should certainly provide excellent train-
ing and considerable enjoyment. The author appropriately closes his
INTRODUCTION by listing a few advanced books to which the inquisi-
tive beginner might turn for more detailed information.

ForDHAM UNIVERSITY JAMES ForBeEs

A Nature Study Book

The Flower Family Album. By Helen Field Fischer and Gretchen Harsh-
barger. 130 pages, 62 full-page plates. The University of Minnesota Press.
1941. $2.50.

When this book first appeared, bound in stiff paper and printed
by the off-set process, it was reviewed briefly in Torreya 40: 212
(1940). It proved so popular that it is now issued by the Univer-
sity of Minnesota Press, bound in attractive cloth covers. The
size remains the same, 81% by 11 inches, with the same attractive
illustrations showing 458 common wild and cultivated flowers
belonging to some forty plant families. The drawings are all to
the same scale, with the height in inches indicated at the side of
the page. For each family a sketch of a single blossom, or flower
cluster is shown to give the family characteristics. Most of these
latter are rather generalized, some without enough detail to be
of much help. In the introduction there is a series of sketches of
flower types with references to the pages where the corresponding
types of flowers are found, this making a sort of key so that one
can hunt more easily to find an unknown plant’s portrait.

Opposite each plate is a description of the family and of the
flowers illustrated. These are ¢ntirely non-technical, often somewhat
whimsical, but clear and accurate. For example the description of
the flowers of the legumes reads, “In most of the family they (the

flowers) look much like butterflies. One petal enlarges to make a
banner to tell the bees that pollen is ready. Two more, called wings,
make a roof landing field for the bees. The last two join to make a
cradle for the ovary, which is wrapped in a gossamer sheet made
from the united stems of the stamens.’ Of the mints we read “The
family is friendly and helpful, seems to love the society of mankind,
for around every dooryard may be found plants of MoTHERWORT
and Catnip, which furnish tonic for man and beast.” Of the com-
posites, “Cooperative Flowers,” “Each disk flower is given the
materials to produce a seed, and it all works out as efficiently as the
production line in an automobile factory.” Of the composites ten
pages of plates include such cultivated forms as dahlia, cosmos,
chrysanthemum, marigold and zinnia and such wild forms as asters,
bidens, dandelion, goldenrod and thistles.

The book will be of little value to the professional botanist, but
the beginning student and gardener will find that a knowledge of the
characters of the plant families given in this informal way will aid
in placing the majority of plants in their proper places while all
plant lovers will find the book helpful and attractive.

GeorGE T. HASTINGS

Butterflies of the North-eastern States

Butterflies. A Handbook of the Butterflies of the United States. Com-
plete for the Region North of the Potomac and Ohio Rivers and East of
the Dakotas. By Ralph W. Macy and Harold H. Shepard. viii + 248 pages.
The University of Minnesota Press. 1941. $3.50.

This attractive book is written for beginners in the study of
butterflies as well as for experienced students and collectors. Cov-
ering all of the north-eastern United States and adjacent Canada
completely it will be useful beyond this area as many, if not most,
of the species described extend beyond any artificial boundary. One
hundred sixty-two species and twenty-seven races are described,
about one-fourth of all the species in North America north of
Mexico. The descriptions include not only the adults but also the
life histories as fully as known, the food plants of the caterpillars,
the ranges throughout the United States and the world—for some,
and as the Mourning Cloak and Red Admiral range over most
of the northern hemisphere. There are often in addition personal

observations on the habits of caterpillars and adults by the authors.
Synonomy is complete and for each species there are page references
to other works in which it is described or illustrated. The keys to
families, genera and species make use of non-technical characters—
wing shape, color and pattern, size—as far as possible so that in
only a few cases is it necessary to consider the venation of the wings
or to use a lens for minute characters. Unfortunately after deter-
mining a specimen by use of the keys no page references are found,
but one must either hunt through the following pages or turn to the
index to find the location of the descriptions.

The book begins with brief accounts of butterflies in folklore,
curious facts about butterflies, protective coloration and mimicry,
sense organs, hibernation and migration, habitats and ranges, classi-
fication and the use of the keys. Half a dozen pages are used in
explaining in detail how to collect, kill and mount specimens, with
suggestions as to making nets, killing jars and display cases.

Twenty-eight butterflies are beautifully illustrated in the four
color plates and there are photographs of thirty-eight others—but
again the lack of page references in the descriptions makes the
locating of the illustrations a matter of search. The book is bound
in green cloth and the press work leaves nothing to be desired.

Both of the authors have been collecting and observing butter-
flies for twenty-five years or more and each has published numer-
ous technical papers and popular articles. Dr. Shepard is Assistant
Professor of Entomology at the University of Minnesota ; Dr. Macy,

Professor of Biology at the College of St. Thomas at St. Paul,

Minnesota.
GeorGE T. HASTINGS

SANTA Monica, CALIF.

Deam’s Flora of Indiana

Flora of Indiana. By Charles C. Deam, M.A., D.Sc., LL.D. With a fore-
word by Stanley Coulter. 1236 pp., with half-tone frontispiece, 2243 distribu-
tion maps and 4 full-page maps in text. Department of Conservation, Indian-
apolis, June, 1940. $3.50. (Obtainable from the State Forester.)

Several reviews of this splendid volume have already been pub-
lished, but the whole story of its excellent features has not been told
yet. The present review seeks to bring out some of its important

points without unduly duplicating what has been said about it
already.

This book is doubtless based on more thorough work than any
other state flora ever published. The author has lived in Indiana
all his life, and has been studying the flora of the state for over
forty years, and he began publishing notes on it in 1904. (He has
also traveled and collected in several other states, and in Central
America.) He had previously published books on the trees, shrubs
and grasses of Indiana, some of them in two or more editions;
and in a long bibliography at the end of the present volume 34 of
the titles, or about 5% of the total, are by him. Good roads and
automobiles, although they have expedited the destruction of natural
vegetation in recent years, have enabled Mr. Deam to visit every
township in the state; something that probably no botanist has
done in any other state.

Accuracy in identification has been his constant aim, and the
aid of several specialists has been enlisted to that end; but of course
there will always be some doubtful cases, on account of intermediate
or imperfect specimens, differences of opinion, or even perhaps
recent mutations in the plants themselves.

In nomenclature the work is very up-to-date. Apparently all
recent revisions involving Indiana plants have been taken advan-
tage of, some as late as 1940 being cited in footnotes. Changes since
the latest manuals available have been surprisingly numerous, and
a great many of the names used will be unfamiliar to readers who
have not kept up with recent developments as closely as Mr. Deam
has. Nearly 22% of the species (not counting varieties and forms)
in his catalogue bear names different from those used in Robinson
& Fernald’s Manual of 1908. Some of these innovations are re-
cently discovered or recently introduced species, some are changes
in classification due to increasing knowledge, and some are due
to differences of opinion as to generic or specific limits, or new
nomenclatorial rules. And Mr. Deam has done very little of the
changing himself, but has accepted the judgment of other workers if
after carefully weighing the evidence he believed it to be valid. A
list of new names appearing for the first time in this book (p. 1112)
includes only 17 cases, and those mostly varieties. In his treatment
of families and genera he has been very conservative, following
Robinson & Fernald’s Manual pretty closely.

35)

In Indiana, as in most other northeastern states, there are few
distinct endemic species as compared with some of the southeastern
states. This is due partly to its small size and dearth of unique
habitats, and partly to the encroachments of civilization, which may
have already wiped out some very local species, and scattered others
outside of their original range. The fact that most of the state was
covered by glaciers, perhaps only 50,000 years ago, may be another
factor tending to reduce the number of endemics. One can pick out
from the catalogue forty or fifty species, varieties, etc., that are at
present known only from Indiana, or Indiana and one other state,
but the great majority of these are hybrids, or recently described
and not very distinct varieties and forms, that might easily turn up
elsewhere when botanists study them closely enough. Practically
none has a well-defined range that stops short of the borders of the
state.

In a state with 94.7 inhabitants per square mile (1940 census),
and the greater part of the area cultivated at one time or another,
and all the forests easily accessible to lumbermen, many unques-
tionably native plants have adapted themselves to changed condi-
tions and persisted in weedy as well as in undisturbed habitats,
while some, less adaptable, or originally confined to sites very
subject to economic exploitation, have disappeared entirely, and
a horde of more or less undesirable immigrants has come in from
Europe and elsewhere to take possession of fields and roadsides.

In Indiana, as in other thickly settled states, practically every
species has felt the devastating effects of civilization in some degree,
and there are all gradations between delicate plants that are found
only in undisturbed habitats, and the weeds of ditches, fields, road-
sides, vacant lots, etc.; so that it is hard to draw the line between
natives and exotics. A few of the species now confined to unnatural
habitats, such as Phytolacca, Prunus angustifolia, Passiflora im-
carnata and Solanum Carolinense, may have existed in Indian clear-
ings before the white man came, but it is hard to get evidence on
that point now.

Many authors of local floras in the northeastern states, with
the veneration for authority characteristic of long-settled regions,
have accepted without question the distinction between native and
introduced species made in current manuals; and if a species is
regarded—trightly or wrongly—as native anywhere in the eastern

United States it becomes ipso facto, in their estimation, native in
the state or county covered by the flora, even if it is there strictly
confined to weedy habitats. Mr. Deam did not go quite to that
extreme, but he gave many weeds the benefit of the doubt, and
classed them as natives. My acquaintance with Indiana vegetation
is chiefly confined to car-window notes in about one-fourth of the
counties, between 1911 and 1941, but from what I know of the same
species elsewhere, I would judge that his 302 introduced species
should be increased to about 500, and the natives correspondingly
reduced.

Many valuable features of the book, such as keys and distribu-
tion maps for every species, the tabular summary, the descriptions
of natural regions, the bibliography of about 700 titles, and the list
of Indiana botanists (142 men and 29 women) have been discussed
by previous reviewers. It is worth noting here that the 41 botanists
who have died lived about 61 years on the average; the later ones
a little longer than the earlier ones.

Typographical errors are very few, and mostly easily detected.
One minor fault of the book is the use of too many fictitious common
names, some of which are longer than the technical names, and not
likely ever to come into general use, and thus serve no useful

purpose. Rotanp M. Harper

UnNIversity, ALA.

RUDD) Tees) Que Aas, CILIUIR
Trip oF AUGUST 23% 1941, To SOUTHERN NEW JERSEY

Mr. Hollis Koster of Green Bank, a competent student of pine
barrens natural history, showed us many interesting species in that
area. Some of the species bring to mind early botanists of the area.
Among these were Rynchospora knieskernu, Lobelia canbyi, Poly-
gala nuttalu, and Panicum commonsianum. The writer was inter-
ested in adding Juncus caesariensis, Snulax walteri, and S. lauri-
folia to the list of plants occurring at the ghost town, Martha.
Mistletoe (Phoradendron flavescens) is a plant that many of us
had not previously seen on a field trip in New Jersey. In the Bass
River State Forest a tree designated as Quercus imbricaria brought

on some discussion of the possibility of its being Q. heterophylla.
As no decision was reached this remains ample reason for a return
field trip.

Mr. Otway Brown guided us over Cape May County and
showed us some of its flora and remarkable plant communities.
The lichenologists worked valiantly this day. Conspicuously large
specimens of a number of trees were brought to our attention. The
climax for most of us was to lay hands on the bald cypress (Taxo-
dium distichum). The specimen is several feet in circumference and
Stone says in 1910 that “very old residents remember them as
being large trees in their youth.” The second tree recorded from
nearby has long since disappeared. We found knees at a distance
of sixty feet or more from the base indicating the extensive root
system. Neither fruit nor seedlings were seen but referring to Stone’s
Plants of Southern New Jersey again we see that at least immature
cones have been observed on this tree. The plant is on the upper
reaches of Sluice Creek in South Dennis. Its natural or introduced
presence is debatable. No lists of species were kept but those col-

lecting found plenty of interest to take. Attendance, seventeen.
Joun A. SMALL

Trip or Aucust 10, 1941—Etystan CLup (Kaiser Roap) To
SUNFISH PoND viA APPALACHIAN TRAIL

From the Elysian Club there was a walk of about a mile over
side trail to the Appalachian Trail. The side trail follows what used
to be a road (Kaiser Road) crossing Kittatiny Ridge from Mt.
Vernon valley to Dimmick Ferry on the Delaware. After picking
up the AT there was a walk of some two miles to Sunfish Pond,
travelling to the southwest. The trail followed a dry ridge at the
start giving fine views of the Delaware valley and the Poconos
beyond. The flora was that of similar portions of Kittatiny Ridge;
oak-hickory with admixtures from the coastal plain such as pitch
pine, scrub oak, and wild indigo. Broom beard-grass were evident
on the exposed outcroppings. Tree species coming into such open-
ings included red cedar, grey birch, poplars, and aspens.

The next mile or so was over richer, more moist terrain, skirt-
ing Tock Swamp for some distance. A larger number of species
were recorded from this portion. Red maple and sour gum were

conspicuous trees while shrubs and ground plants were represented
by numerous species. Four northern plants were seen which venture
into New Jersey only in these upland areas of the northern coun-
ties. Thus New Jersey represents the southern limit of their range
except as they follow down the higher ridges of the Appalachians.
Cornus canadensis is one of these. Neither Britton’s Catalog nor
Taylor’s Flora of the vicinity of New York record this species
from Warren County. A specimen in the herbarium of the New
York Botanical Garden is labelled, “Green’s Pond, Warren County,
New Jersey, May 21, 1921. In larch woods, very rare.” Green’s
Pond: is now Mountain Lake on the topographical maps. Rhodo-
dendron canadense is reported in the above manuals from Morris
and Sussex Counties only. Prunus cuneata is also limited to the
northern counties in New Jersey, and Muhlenbergia racemosa is
credited with a similar distribution. The last three are not recorded
from Warren County in the herbarium of the New York Botanical
Garden. This trip therefore may have produced three definite exten-
sions of range.

Among the ferns seen, Aspidium simulatum and Woodwardia
virginica are distributed over the state but their occurrence is
sufficiently local to make this station of interest. A total of ten
ferns and 138 flowering plants were recorded without leaving the
trail. The list is filed with the field committee. Attendance, ten.
Leader, L. Hardy. Plant lists by L. E. Hand and G. G. Nearing.

Joun A. SMALL

Trip oF Octoser 4, 1941, ro BRookKLYN BoTANIC GARDEN

This walk was devoted to a study of the pines. The distinguish-
ing characteristics of the white pines and pitch pines were pointed
out. The circumboreal distribution of the genus was evident from the
walk. American species included our common white pine, Pinus
Strobus, and the dominant pitchpine of southern New Jersey, P.
rigida. Going northward on the magic needles we saw the red pine,
P. resinosa, and the far northern jack pine, P. Banksiana. Moving
westward via the Allegheny species, P. pungens, we noted the west-
ern P. flexilis. It was observed that two of the western species, P.
ponderosa and P. Jeffreyi do not grow well in the Prospect Park
environment. The following European species were seen: P. nigra

and P. sylvestris which are not uncommon in American nurseries,
then P. Heldreichu, P. mugo and P. Cembra. The Himalayan pine,
P. excelsa, was the largest specimen among Asiatic pines. Other
species were P. Thunbergu, which might be called silver-bud pine,
P. densiflora, P. parviflora, P. koraiensis and P. Bungeana which
grows as a shrub here. The leader stated that the genus Pinus
reaches into the southern hemisphere in only one place, the moun-
tains of Java. Has anybody been looking at Java on the map lately?
Attendance was 11. Leader: Dr. Alfred Gundersen.

Joun A. SMALL

TRIP OF OCTOBER 18 To THE BROOKLYN BOTANIC GARDEN

This time the objective was the study of evolution. The Con-
servatory was visited and we were shown the exhibit of the principal
groups of plants. Algae, Ferns and Angiosperms are the three main
stages of plant evolution, with many diverging lines within and be-
tween. Algae are nearly all water plants. Early stages of land plants
are suggested by lichens, by liverworts, by the fossil Rhynia group,
and also by clubmosses and horsetails. These have no true leaves,
but fronds of ferns are primitive leaves, which are now recognized
as fused and flattened branches. Seeds differ from spores somewhat
as the large eggs of reptiles differ from the small eggs of amphibians,
for seeds and reptile eggs are adaptations to land life. Differences
such as open or closed ovary should be considered with the time
difference between Gymnosperms (late Devonian) and Angiosperms
(Cretaceous). The significant characteristic of flowering plants is
insect pollination, although a few of them have reverted to the primi-
tive wind pollination. Deciduous leaves and the herbaceous habit
must be thought of as adaptations to a winter season. Various books
were seen and discussion closed the meeting. Attendance 17.
Leader: Dr. Alfred Gundersen. There will be other visits for some-

what different studies in 1942.
Joun A. SMALL

Trip oF NoveMBER 16, 1941. Kaiser Roap To MILLBROOK Roap
Via APPALACHIAN TRAIL, WARREN County, N. J.

Like the trip of August 10, this walk started from the Elysian
Club, following the side trail to the crest of the ridge. Several plants

of the stiff stemmed gentian were seen along the trail up the moun-
tain. The AT to the southwest leaves Kaiser Road at the crest of
the ridge. The road then leads northward crossing the ridge in an
apparent quest of a suitable line of descent. When this point is
reached the AT proceeds northeast following the skyline. And a
veritable skyline it is: narrow, with occasional high outcroppings
that make delightful natural rock gardens. In these one finds Aqui-
legia, mountain phlox, marginal shield fern, polypody, and a variety
of mosses and lichens. The view from these points is excellent, par-
ticularly at this season of the year.

At other points the trail follows the top of an escarpment drop-
ping from the narrow summit ridge to the valley below. One sees
Andropogon and other grasses in such places, with rock tripes on the
stone. Red cedar, sweet fern, and blueberries make up the major por-
tion of the woody flora. The general flora of the ridge is oak, of
which eight species were recorded, among them Q. prinoides which
we note is not credited to Warren County in Britton’s survey.
Hickories, beech, red maple, black birch, ash, and sour gum made up
the other common tree species. Flowering dogwood, azaleas, and
laurels indicated the beauty to be seen along this trail at the appro-
priate season.

About three miles above Kaiser Road the trail descends abruptly
to a notch in the ridge where another road, now long abandoned,
formerly crossed to the Pahaquarry copper mines. This road is still
maintained eastward as an access road to the nearby boy scout camp
which owns most of the land over which we had travelled. This
makes a suitable point for approaching the trail by car. Several spe-
cies were added along the brook at the point where we crossed the
notch. We then climbed abruptly as the ridge regains its normally
rather level crest. Along the trail at this point, a well-fruited sprout
growth of Castanea was found. None of the fruits examined ap-
peared viable however.

The vegetation continued as on the south side of the notch. Even
more spectacular escarpments were encountered but with more
limited views. Time and daylight did not permit us to continue to
Millbrook road, so a side trail to the west was taken, bringing us
around to Catfish pond. From here we crossed the scout camp and
went out over their road. Attendance 11. Leaders, Mr. and Mrs.
Louis Anderson. Plant lists by Louis Hand and W. L. Dix. These

lists recorded 111 flowering plants, 7 ferns, 8 mosses, 20 lichens,
and 3 fungi. The lists for mosses and fungi are only fragmentary
since no one well qualified to list these plants was available. One of
the lichen species, Cladonia turgida was remarkable for being the
second collection in the state, the other being from Atlantic County
where it was collected in 1936 by the late Raymond H. Torrey.

Joun A. SMALL

HvO CED DEINGSTOE DEE GEUB
MINUTES OF THE MEETING ON JANUARY 6, 1942

The annual meeting of the Torrey Botanical Club was held at
the Men’s Faculty Club of Columbia University on Tuesday, Janu-
ary 6, 1942. Dinner was served at 6:30 p.m., after which the meet-
ing was called to order by the President, Dr. J. S. Karling. Seventy-
seven members and friends were present.

The minutes of the previous meeting were approved as read.

Reports of the officers of the Club were mimeographed and dis-
tributed to those present at the annual dinner. The report of Dr.
Dodge as delegate to the Fordham University anniversary and to
the A. A. A. S. meetings in Dallas, and the report of Dr. Robbins
as delegate of the Club to the New York Academy of Science were
accepted by the Club.

Dr. Small announced that as a member of the New York-New
Jersey Trail Conference, the Club was participating in the Sports-
man’s Show.

The Club accepted the 1942 budget as approved by the Council.

Dr. Zimmerman moved that the Club reconsider its action of
May 16, 1941, dedicating the 1942 volume of the Bulletin to Dr.
Harper, and extend the dedication to include the surviving charter
member of the Club, Dr. Denslow. Dr. Bold seconded the motion
and the Club so voted.

The President announced that the following list of officers had
been elected by the Club to serve during 1942:

President: C. Stuart Gager
First Vice-President : John A. Small

Second Vice-President : Clyde Chandler
Corresponding Secretary: H. C. Bold

Recording Secretary : John W. Thomson, Jr.
Treasurer: W. Gordon Whaley
Editor : Harold W. Rickett
Bibliographer : Lazella Schwarten
Business Manager: Michael Levine
Members of the Council: John M. Arthur
Lela V. Barton
Arthur H. Graves
Edwin B. Matzke
Delegate to the Council of the New York Academy of Sciences:
W. J. Robbins
Representative on the Board of Managers of the New York Botanical
Garden: Henry A. Gleason
Representatives on the Council of the American Association for the
Advancement of Science: Albert E. Hitchcock
John S. Karling

Mr. Rutherford Platt then conducted a guessing game with the
Kodachrome colored slides of plants for which he is so well known.
The botanists present did not prove to be too familiar with the com-
mon plants which he showed. First, second and booby prizes were
awarded.

The meeting was adjourned at 9:20 P.M.

Respectfully submitted,

JoHN W. THOMSON, JpR.,
RECORDING SECRETARY

MINUTES OF THE MEETING OF JANUARY 21, 1942

The meeting was called to order at 3:35 p.m., in the Member’s
Room of the New York Botanical Garden by the President, Dr. C.
Stuart Gager. Thirty-six members and friends were present. In the
absence of a Recording Secretary, the Corresponding Secretary read
the minutes of the preceding meeting. These were adopted as read.
The following were elected unanimously to annual membership in
the Club:

Dr. W. B. Baker, Emory University, Atlanta, Georgia
Mr. Stanley D. Wikoff, 91 Easton Avenue, New Brunswick, N. J.

Mr. G. Thomas Robbins, University of Colorado, Boulder, Col.
Mr. Frank G. Lier, 510 West 110th Street, New York City.

The transfer of Mr. W. Herbert Dole, 25 Overlook Avenue, West
Orange, N. J., from annual to associate membership was approved.
The following resignations were noted with regret:

From annual membership:

Mr. H. H. McKinney, Horticultural Field Station, Beltsville, Md.
Mr. Alan Martin, Glenwood, N. J.

Prof. A. J. Sharp, University of Tennessee, Knoxville, Tenn.

Miss Olga H. Hingsburg, 46 Esplanade, Mt. Vernon, N. Y.

From associate membership:

Mr. Seymour Barrett, 1475 Grand Concourse, N. Y. C.

Dr. George H. Hallett, Jr., 3353 82nd Street, Jackson Heights, N. Y.
Mrs. Ruth D. Hallett, 3353 82nd Street, Jackson Heights, N. Y.

Mr. L. W. Steiger, 835 Summit Avenue, Hackensack, N. J.

The Corresponding Secretary announced that the Council had
accepted the resignation of Dr. Thomson as Recording Secretary,
and read his letter of resignation to the Club. The President stated
that in accordance with the Constitution the Council had elected
Miss Honor Margaret Hollinghurst to fill the unexpired term of
Recording Secretary.

The President also announced that the Council had sent a tele-
gram of congratulations to Professor R. A. Harper in the name of
the Club, this being Dr. Harper’s 80th birthday.

The President also announced that through an error in the bal-
lot, a vacancy existed in the Council membership. Dr. Rickett nomi-
nated Dr. W. J. Bonisteel for the vacancy, the nomination was sec-
onded by Dr. Robbins, and Dr. Bonisteel was unanimously elected
to a term from 1942-1944.

Dr. Small again announced that tickets were available for the
Sportsman’s Show.

The scientific portion of the program consisted of a report by
Dr. L. V. Barton of the Boyce Thompson Institute on “Some Spe-
cial Problems in Seed Dormancy.” The speaker’s abstract follows:

Dormancy in relation to seeds is a general term used to indicate the failure
of the embryo to resume growth when placed under conditions of temperature,
moisture, and oxygen supply which ordinarily bring about germination. The
dormant state may be imposed by seed coats, dormant embryos or a combina-
tion of seed coat and dormant embryos. Furthermore, there are seeds in which
the root is not dormant but the shoot or the bud which forms it is dormant.
In the last case it is necessary to treat for a period at a low temperature (1°
to 10° C.) in a moist medium after the root has already formed in order to
break the epicotyl dormancy so that the first green leaf may develop. Such
treatment may be given effectively at any time between the first appearance of

the radicle and the maximum development of the root system from the stored
food in the seed.

Recent experiments in which seeds of Convallaria majalis L. and Smilacina
racemosa (L.) Desf. were the test material, showed epicotyl dormancy of a
different type in that the period at low temperature, in order to be effective,
must be given, not merely after root production, but after the seedlings had
developed to the stage where their shoots had broken through the first enclos-
ing sheaths. Exposure at earlier developmental stages was without effect in
breaking epicotyl dormancy. Three to five months at 5° or 10° C. was found
to be necessary for forcing the first green leaves of Convallaria and Smilacina.

Low-temperature pretreatment of the imbibed seeds increased root produc-
tion in Convallaria and was essential to root formation in Smuilacina when
plantings were made in the soil in the greenhouse.

After considerable discussion, the meeting adjourned at 4:40

p.m., to enjoy tea and other refreshments served by members of the
Garden Staff.
Respectfully submitted,

Harotp C. Bo rp,
Acting Re. Sec.

NEWS NOTES

The Torrey Club has undertaken a botanical survey of that
portion of the Appalachian Trail that is maintained by the New
York-New Jersey Trail Conference, namely, from the Delaware
River to the Connecticut state line. Something over twenty miles
of trail and alternate trail between the Delaware and Flatbrooks-
ville road were covered last season. Three hundred and sixty-seven
species of Spermatophytes, 28 species of Pteridophytes, 24 species
of Bryophytes, 105 species of lichens (disregarding forms, modes,
and varieties), and 47 species of fungi have been recorded to date.
The only alga so far determined is Microspora stagnorum. In
many of the groups a considerable number of additional species
would be recorded if specialists in those plants were available on
the trips. The project will be continued this season.

THE TORREY BOTANICAL CLUB
OFFICERS FOR 1942

President: C. Stuart GAGER
Treasurer: W. Gorpon WHALEY
Editor: Harotp W. RIcKETT

Vice-Presidents: JoHN A. SMALL, F,
CLyDE CHANDLER

Recording Secretary: Miss Honor M.

HoLLINGHURST Business Manager: MicHarL LrEvIne
Corresponding Secretary: Harotp C. Bibliographer: Mrs. LazELLA SCHWAR-
Bop TEN

Delegate to the Council, N. Y. Academy of Sciences: W. J. Ropstns

Representatives on the Council of the American Association for the
Advancement of Science:

A. E. HitcHcock J. S. Karrine

Representative on the Board of Managers of the N. Y. Botanical Garden:
H. A. GLEASON

Council for 1940

Ex officio members
Edwin B. Matzke
Harold N. Moldenke

John S. Karling
Florence C. Chandler

Michael Levine
William J. Robbins
Henry K. Svenson
John A. Small

Bernard O. Dodge
Arthur H. Graves
Alfred Gundersen
George T. Hastings
Harold W. Rickett

Elected members

1940-1942
Lela V. Barton
Ralph H. Cheney
Robert A. Harper
Edmund W. Sinnott

1941-1943
Helen M. Trelease
Ralph C. Benedict
John H. Barnhart

1939-1941
Gladys P. Anderson
John M. Arthur
Harold H. Clum
Percy W. Zimmerman

Committees for 1940
ENDOWMENT COMMITTEE

Clarence Lewis, Chairman J. Ashton Allis
Caroline C. Haynes Henry de la Montagne Helen M. Trelease

ProGRamM COMMITTEE

Harold C. Bold, Chairman (e- elicte)
William J. Robbins
E. B. Matzke

FIELD COMMITTEE

John A. Small, Chairman
Rutherford Platt
Henry K. Svenson
Ellys Butler
Dolores Fay
Eleanor Friend

Edward J. Alexander
G. G. Nearing
Vernon L. Frazee
Alfred Gundersen
Inez M. Haring

H. N. Moldenke

Locat FLora COMMITTEE

W. H. Camp, Chairman
Harold W. Rickett
Ora B. Smith
Herbert M. Denslow

William J. Bonisteel
James Edwards
John M. Fogg, Jr.

Cryptogams

E. Marcy
P. W. Zimmerman
G. Whaley

Robert Hagelstein
Michael Levine
James Murphy
Daniel Smiley, Jr.
Farida A. Wiley

Dolores Fay
H. Allan Gleason
Hester M. Rusk

Ferns and Fern Allies: R.C. Benedict, W. Herbert Dole, N. E. Pfeiffer

Mosses: E. B. Bartram

Liverworts: A. W. Evans, E. B. Matzke
Freshwater Algae: H.C. Bold

Marine Algae: J. J. Copeland _

Fungi: A. H. Graves, J. S. Karling, W. S. Thomas
Lichens: J. W. Thomson, Jr.

Myxomycetes: R. Hagelstein

CoMMITTEE ON EXCHANGES

Harold C. Bold Amy Hepburn

Elizabeth Hall

OTHER PUBLICATIONS
OF THE

TORREY BOTANICAL CLUB

(1) BULLETIN

(2) MEMOIRS

Volume 18, no. 1, 108 pages, 1931, price $2.00. Volume 18, no.
2, 220 pages, 1932, price $4.00. Volume 18 complete, price $5.00.

Volume 19, no. 1, 92 pages, 1937, price $1.50. Volume 19, no.
2, 178 pages, 1938, price $2.00.

(3) INDEX TO AMERICAN BOTANICAL
LITERATURE

Reprinted monthly on cards, and furnished to subscribers at three
cents a card.
Correspondence relating to the above publications should be
addressed to
W. Gordon WHALEY,
Barnard College,
Columbia University,
New York, N. Y.

Herbarium,

Volume 42 May-June, 1942 Number 3

TORREYA

A Bi-MonTHLy JoURNAL oF BoTANICAL NoTEs anp NEws
. EDITED FOR
THE TORREY BOTANICAL CLUB

WILLIAM J. BONISTEEL

John Torrey, 1796-1873

CONTENTS

Program of the Torrey Botanical Club—The Seventy-fifth Anniversary Cele-

pation tese statis we ice CE Ne ce sae etn ae ar toon eos Guede he arn MIR dey igen tet SES
Chile Tarweed in Quebec.......................00005. Haroitp N. MoLtpENKE
Collecting Chicle in the American Tropics (Part 2)......... JoHN S. KARLING
More Fungi from the Front Lawn.......................... Laura A. KoLk
Book Reviews

AboutO@urselvesi/. hee Rh ee LAr ee MIcHAEL LEVINE

Microbe’s Challenge... 0.0.0... .00.0. 00 0c e ec eee eee RuHopa W. BENHAM

Practical Plant Anatomy............................ W. Gorpon WHALEY

Fundamentals of Plant Science........................ Epwin B. MatzkrE
velde Drips softs the x@lubron ye a5 oe Nn WEN Pe ce ea oy See a

An Herbal (1525)....... Bal cla ae SER PEM ET Bin Hu hers Get H. W. RicKeEtTT

Proceedings of the: Clubs g hep ey Rae REN Shes cue aan
News i NOtesi sige steele ns aie SU a eae I SOL Se CMe oll te oer pele

PUBLISHED FOR THE CLUB

By THE FREE PRESS PRINTING COMPANY
187 COLLEGE STREET, BURLINGTON, VERMONT

Entered as second class matter at the post office at Burlington, Vermont,
October 14, 1939, under the Act of March 3, 1879

TORREYA

TorrEYA, the bi-monthly publication of the Torrey Botanical Club, was established
in 1901. TorREYA was established as a means of publishing shorter papers and inter-
esting notes on the local flora range of the club. The proceedings of the club, book
reviews, field trips and news notes are published from time to time. The pages of
TORREYA are open to members of the club and others who may have short articles
for publication.

Claims for missing numbers should be made within sixty days callenniee their
date of mailing. Missing numbers will be supplied free only when they have been
lost in the mails. All subscriptions and requests for back numbers should be ad-
dressed to the treasurer, Dr. W. Gordon Whaley, Barnard College, Columbia Uni-
versity, New York, N. Y.

Of the annual membership dues of the Torrey Botanical Club, $.50 is for a year’s
subscription to TORREYA.

INSTRUCTIONS TO CONTRIBUTORS

The manuscript should be prepared so that it conforms to the best practice as
illustrated by current numbers of TorreyA. Manuscript should be typed double-
spaced on one side of standard paper. The editors may accept papers up to eight
printed pages in length. Longer papers may be published if the author agrees to bear
the cost of the additional pages. Illustrations (including tables and graphs) should
not exceed twelve per cent of the text; authors of more copiously illustrated ar-
ticles may be asked to pay for the excess material. Brief notes will be published
with especial promptness.

TorREYA is edited for the Torrey Botanical Club by

WM. J. BONISTEEL
W. H. CAMP DOROTHY J. LONGACRE

MEMBERSHIP IN THE TORREY BOTANICAL CLUB

Manuscripts for publication, books and papers for review, reports of field
trips and miscellaneous news items should be addressed to:

DR. WM. J. BONISTEEL
BIoLoGIcAL LABORATORIES, FoRDHAM UNIVERSITY
New York, N. Y.

TORREYA

VoL. 42 May-JUNE No. 3

The Torrey Botanical Club
Seventy-fifth Anniversary Celebration
June 22 to June 27, 1942

Monday, June 22.

10:00 a.m. to 12:30 p.m. Registration. Rotunda, Low Memo-
rial Library, Columbia University—General informa-
tion about accommodations in Johnson Hall, Livingston
Hall, and King’s Crown Hotel available at registration.

2:00 to 4:30 p.m. Scientific Program. Room 305 Schermerhorn

Hall. Sectional chairman: Dr. Edwin B. Matzke.

“The History of Botany at Columbia University.’ Dr.

John S. Karling.

Symposium on Morphology.

1. “Haphazard as a Factor in the Production of Tetra-
kaidecahedra.” Dr. F. T. Lewis, Harvard Medi-
cal School, Cambridge, Mass.

2. “The Evolution and Determination of Sexual Char-
acters in the Angiosperm Sporophyte.” Dr. C. E.
Allen, University of Wisconsin, Madison, Wis.

3. “The Leaf-Stem Relationship in Vascular Plants.”
Dr. R. H. Wetmore, Harvard University, Cam-
bridge, Mass.

4. “Problems of Pattern in Plant Development.” Dr.
E. W. Sinnott, Yale University, New Haven,
Conn.

4:30 to 6:00 p.m. Tea and Torrey Exhibit. Rotunda, Low
Memorial Library.

7:00 p.m. Anniversary Banquet. Men’s Faculty Club, 117th
St. and Morningside Drive.

Presentation of President of the Torrey Botanical Club,

Dies CC, Swat (Gees ose esos acs Dr. John S. Karling

Presentation of officially appointed delegates
Dr. John S. Karling

TorreYA for May-June (Vol. 42, 65-103) was issued June 5, 1942.
65

Welcome to delegates, members, and guests

Pres. C. Stuart Gager
Response of delegates
Reading of letters of felicitation......... President Gager

Tuesday, June 23.

10:00 a.m. to 12.30 p.m. New York Botanical Garden. Sec-
tional chairman: Dr. Wm. J. Robbins.

“The History of the New York Botanical Garden.” Dr.

Wm. J. Robbins, Director.

Symposium on Taxonomy.

1. “Contributions of the Torrey Botanical Club to the
Development of Taxonomy. Dr. H. A. Gleason.
New York Botanical Garden.

“Modern Taxonomy and Its Relation to Geography.”

Dr. H. K. Svenson, Brooklyn Botanic Garden.

3. “Economic Aspects of Taxonomy.” Dr. E. D. Mer-

rill, Harvard University. |

4. “The Importance of Taxonomic Studies of the

Fungi.” Dr. F. D. Kern, Pennsylvania State
College, State College, Pa.

12:30 to 2:00 pm. Basket Luncheon in the Rock Garden,
50 cents.

2:00 to 4:30 p.m. Inspection tour of Gardens, Conservatories,
Laboratories, and Herbarium, conducted by members of
the Staff.

8:30 p.m. Smoker. Men’s Faculty Club, Columbia University.

Wednesday, June 24.

10:00 a.m. to 12:30 p.m. Scientific Program. Boyce Thompson
Institute for Plant Research, Yonkers, New York.
Sectional chairman: Dr. P. W. Zimmerman.

“The History and Organization of the Boyce Thompson

Institute. Dr. Wm. J. Crocker, Director.

Symposium on Growth.

1. “Viruses in Relation to the Growth of Plants.”
Dr. L. O. Kunkel, Rockefeller Institute for Medi-
cal Research, Princeton, New Jersey.

2. “Morphogenetic Influences of Plant Hormones.” Dr.
P. W. Zimmerman, Boyce Thompson Institute.

3. “The Many-sided Effects of Animal Hormones and
Their Possible Resemblance to Plant Hormones.”
Dr. Oscar Riddle, Carnegie Institute of Wash-
ington, Cold Spring Harbor, Long Island, New
York.
12:30 to 2:00 p.m. Luncheon. Boyce Thompson Institute acting
as host.
2:00 to 4:30 p.m. Inspection tours of grounds and laboratories
conducted by members of the Staff.
Exhibits by investigators of the Institute.
8:30 p.m. Public Lecture. American Museum of Natural
History. “Plants Need Vitamins, Too.” Dr. Wm. J.
Robbins, New York Botanical Garden.

Thursday, June 25.
10 :00 a.m. to 12:30 p.m. Scientific Program. Brooklyn Botanic

Garden, 1000 Washington Avenue, Brooklyn, New

York.

Sectional chairman: Dr. C. Stuart Gager.

“The History of the Brooklyn Botanic Garden.” Dr. C.

Stuart Gager, Director.

Symposium on Genetics.

1. ‘Genetics, the Unifying Science in Biology.” Dr.
George H. Shull, Princeton University, Prince-
ton, New Jersey.

2. “A Consideration of Criteria of Center of Origin.”
Dr. Stanley Cain, University of Tennessee,
Knoxville, Tennessee.

3. “The Status of Plant Pathology in 1875 and in
1942.” Dr. George M. Reed, Brooklyn Botanic
Garden.

4. “Technical Applications of Genetics in Plant Breed-
ing in 75 Years.” Dr. A. F. Blakeslee, Carnegie
Institution of Washington, Cold Spring Harbor,
Long Island, New York.

12:30 to 4:30 pm. Luncheon. Brooklyn Botanic Garden. 50
cents. Inspection tours of gardens and laboratories con-
ducted by members of the Staff.

Friday and Saturday, June 26 and 27.

Two-day field trip to Southern New Jersey. Dr. John A. Small,

New Jersey College for Women, Field Chairman.

First day to Seaside Park for beach, salt marsh, and en-
croaching pine barren vegetation. Overnight accommodations
at Toms River.

Second day to the dry barrens, “The Plains,” and the bogs.

Chile Tarweed in Quebec

Harotp N. MoLpENKE

Since the publication of my recent note on the occurrence of Chile
tarweed east of the Mississippi River (Torreyva 41: 162-164), my
good friend, Brother Marie-Victorin, of the Montreal Botanical
Garden, has kindly sent me some more material of this species, rep-
resenting the first known eastern Canadian records. All these speci-
mens appear to be the typical form of Madia sativa Molina, rather
than the variety, and all except three from the Marie-Victorin
herbarium are deposited in the herbarium of the Montreal Botanical
Garden.

The first specimen is an undated one, collected by Omer Caron
in Lotbiniere County, Quebec. The earliest dated collection is rep-
resented by five sheets (two in the Montreal Botanical Garden her-
barium and three in the Marie-Victorin herbarium) collected by
Brothers Marie-Victorin and Rolland-Germain on August 24, 1927,
in uncultivated ground along the road from Longueuil to Gentilly,
Chambly County, Quebec (no. 29062), where the collectors state
that the species was introduced and abundant. On September 16,
1933, the same two collectors found it naturalized in fields at
Longueuil (no. 45645, two sheets). On August 20, 1935, the same
collectors collected it again in an abandoned field at Longueuil (vo.
43637, two sheets), and on September 14, 1935, Cécile Lanouette .
collected it along Chemin du Lac at Longueuil, where it seems, there-
fore, to be very definitely established.

New York BotTANICAL GARDEN

Collecting Chicle in the American Tropics
(Part 2)

JoHN S. KARLING

IDENTIFICATION OF ACHRAS SPECIES AND CHICLE ADULTERANTS

In spite of the economic importance of chicle and the sapodilla
tree, there is still some confusion and ignorance among contractors,
chicleros, and professional botanists about the sources of chicle and
the substitutes commonly used. A large number of species of differ-
ent families yield gum which is utilized to a limited extent in chew-
ing gum manufacture, but there is little doubt that the best and larg-
est supply of chicle comes from Achras zapota, although some tax-
onomists have denied this. This species as described by Plumier
(1703), Linnaeus (1753, 1762), Jacquin (1760, 1763), Brown
(1789), Pierre and Urban (1904), Coville (1905), Cook (1913),
Pittier (1914, 1919), Hummel (1925), and Standley (1925, 1932)
appears to be quite variable, and confusion as to the source of chicle
is to be expected, especially when the herbarium material has been
collected under different vernacular names from widely separated
localities. In 1888 Planchon listed three species of Achras as com-
mercially important, and recently (1919) Pittier added two addi-
tional latex-yielding species, A. chicle and A. calcicola, which were
formerly included in A. zapota. Pierre and Urban (1904) described
four varieties of A. zapota on the basis of fruit and flower sizes and
shapes. Whether or not these latter are valid species is uncertain,
but it is not improbable that when the jungles of southern Mexico,
Central and South America have been thoroughly combed and the
forms carefully studied, additional species and varieties will be seg-
regated.

In British Honduras the native chicleros, according to Hummel
(1925), recognize the following types of A. zapota:

(1) “Female Sapodilla’—by far the best tree for producing
chicle. Large edible fruit of good quality. Leaves smaller and closer
together than those of any of the other kinds of sapodilla. However,
the leaves of saplings are often abnormally large and their size and
shape are, therefore, misleading. This tree is more numerous in the
north of the Colony than in the south ; the Sibun River may, roughly,
be taken as the dividing line betwéen good and inferior chicle.

“Female sapodilla”’ grows well on inferior soil, on so-called “Broken
Ridge” soil, but it grows also on the best soil together with ma-
hogany.

(2) “Crown Sapodilla’—produces the second best chicle. The
general appearance of this tree is so similar to No. (1), “female
sapodilla,” that even chicleros are not always certain in distinguish-
ing it, unless they can see the fruit, which is much smaller and of a
slightly different and more elongated shape from those of the “female
sapodilla,”’ and not quite so delicious to eat as the latter.

(3) “Male” or “Bastard Sapodilla’—produces little chicle, less
fluid and of inferior quality. The leaves are considerably larger and
further apart than those of Nos. (1) and (2). The fruit is small,
inedible and grows in small bunches. This tree does not bear fruit
every year. The belief that it does not bear any fruit at all is wide-
spread. The attribute “male” has no botanical significance ; it 1s ap-
plied in the native nomenclature quite generally to plants of inferior
quality, while the attribute “female” is here usually used for plants
of superior quality.

(4) “Chicle Bull’—the most useless of the various sapodilla
trees. The leaves are smaller than those of the “male sapodilla.” It
is usually recognized by its fruit, which are the size of grapes and
grow in fairly great bunches almost like grapes.

The “male” sapodilla tree or “chicle macho” (Record and Kuy-
len, 1926) of British Honduras, as reported by Hummel, has been
described from Guatemala and Mexico, where Pittier treated it as a
new species, 4. chicle. On the basis of reports which he received
from chicleros, Pittier regarded this species as the chief source of
chicle, and stated: “The chicle of commerce is not extracted exclu-
sively, if at all, from the latter species, Achras zapota.” This conten-
tion has been severely criticized and is undoubtedly wrong, or at
least certainly needs additional proof. Record and Kuylen report that
the latex of A. chicle is used only to a limited extent for chicle. Hum-
mel’s descriptions of “chicle bull” and “male” sapodilla are the same ~
with respect to size and growth of the fruits in bunches. This claim
has also proven to be incorrect in most instances, since “chicle bull”
in the crown lands of British Honduras is very similar to “female”
sapodilla with respect to fruit, etc. However, in 1927 Record re-
ported 4. chicle from Honduras with large edible fruits, which indi-

cates further the variability of Achras species and the difficulty of
distinct differentiation. |

The confusion about the species of Achras which yield the chicle
of commerce stems largely from the fact that the Indians and chic-
leros in various localities and countries have different names for
the same plant or the same name for widely different plants; and
collectors unfamiliar with these vernacular synonyms may be readily
led astray. Use of vernacular names as a criterion of differentiation
without excellent herbarium material is worthless and leads at once
into difficulties. To illustrate, Achras chicle in British Honduras is
generally known as “chicle macho” and often as “chicozapote,’ while
in Guatemala, according to Record (1926), it goes under the name
of “nispero” or “zapotillo.” “Chico zapote”’ and “zapotillo,’ how-
ever, are two of the vernacular names generally applied to Achras
zapota in Mexico and other regions. Similarly, “sapodilla” is applied
to A. calcicola in Panama. Hence, chicle reported to come from
“chico zapote,” “‘zapotilla,” and sapodilla may involve several species
of Achras as well as other genera of the Sapotaceae.

Throughout Mexico, Central America, South America, and the
West Indies, Achras zapota has more than twenty different vernacu-
lar names, many of which are also applied to trees of entirely differ-
ent species and genera. Below is a partial list of names commonly
given to Achras zapota in different parts of tropical America, accord-
ing to Pittier (1914), Standley, and others.

West Indies, Venezuela, Colombia, and
generally throughout Central Amer-

Ged tinea eee neta leu lee eenay oan ASN Ps Nispero zapote

British West Indies and Florida...... Naseberry, Sapodilla, neesberry, nis-
berry

French West Indiés................. Sapotier, sapotille, sapotillier

Dutch West Indies................... Mispel, mispelloom

Guatemala ty yactiee dh tee een aoktte Chicle zapote, muy, chico zapote, sapo-
dilla, zapote chico, zapotillo

RstIGa talline neers cust dendiate uc cneeumle ts Zapote, ya, zapote de abejas, tzapotl,
palo maria, zapotillo, peruetana

Were, Cir, OAS8iCs soncocodcov0cdu0: Zapote, chico zapote, zapote chico,
chico, zapotillo, guendaxina, txicoza-
potl

Panama ..... LeS5S SHG Ao MOOT ORDO SOOM Mamey, zapote

Sailkievolore vs ee tcev amin amin eee neeee enc ate Muyozapot

GostamRicaae mea crns ceeere tes unison Korok, zapote, zapotillo

INiCarag tains: Shy toeiees atoms cme tare ntevete Zapote, iban, zapotillo, chico.

ElonduraSmee ee seen eee eee soe ee NispeLrouzapoteszaporlllo
Bicuadomerracnaceer ae hie oo eeoe er NISperolrguite;nse
Brazile escear ss ct ann encase Sapote, sapotilla

The terms “nispero,’ “zapote,’ “zapotillo,’ used generally
throughout Central America for A. zapota, are often extended to
include several genera and species of the Sapotaceae, such as Sider-
oxylon amygdalinum, S. Gaumeri, S. Meyeri, Lucuma salicifolia,
L. Durlandu, Dipholis Stevensonu, Calocarpum mammosum, C.
viride, Chrysophyllum oliviforme, and others. It is not surprising
then that in the early attempts to classify the chicle-yielding trees
confusion arose among collectors in separate localities. In recent
years, however, more extensive botanical collections have been made
in the chicle areas, and the commercially important trees are fairly
well known.

Achras zapota itself, as noted before, shows considerable varia-
tion in different localities, and a number of local varieties are recog-
nized by the chicleros. In regions south of the Belize River, British
Honduras, and in certain localities in Peten, Guatemala, is found the
form of A. gapota which is generally known as “chicle bull” or
“chiquibul.” Taxonomically and morphologically, as far as it has
been studied, it is reported to be the same as A. zapota, but for
chewing purposes its gum is very inferior to that of trees growing
north of this river. The latex is difficult to coagulate and requires
longer boiling, while the resulting gum must be worked and fre-
quently washed before it can be molded and hardened sufficiently for
shipment. Since trees yielding “chicle bull’? are commonly found on
a slightly different type of soil, the difference in quality of the gum
has been attributed to this variation. The sapodilla tree appears to
flourish best on calcareous marl and disintegrated. limestone which
predominate in the Yucatecan Peninsula of Mexico, northern British
Honduras, and the Peten District of Guatemala ; and it is primarily
from this contiguous area that the best A. zapota chicle comes.
South of this region the surface soil is reported to be less limey, and
here occur A. chicle, “chicle bull,” and the so-called “bastard sapodil-
las”
chicle operators as necessary for good chicle. It is not uncommon,
however, in regions where they overlap to find the two kinds of
sapodillas growing side by side in the same type of soil, but still
showing a marked difference in gum quality. In view of this it is

in greater abundance. A soil rich in lime is thus regarded by

not improbable that “‘chicle bull” may be a variety or physiologically
differentiated race of A. zapota.

Within the species A. zapota, which yields the best chicle of com-
merce, most chicleros in British Honduras recognize three forms:
zapote blanco, zapote colorado, and zapote morado. The mauve or
morado is said to be the best yielder, with the white next in order.
The white and red forms are also recognized in Mexico and Guate-
mala, but the mauve is not generally distinguished. In those coun-
tries the white sapodilla is reported to yield almost twice as much as
the red (Anonymous, 1923). Whether or not these three forms also
are to be recognized as varieties or physiological races of A. zapota
remains to be seen. Morphologically they appear the same. Their
difference lies chiefly in the color of the bark, and the distinctions
may be so fine that they are often unrecognizable except to the prac-
tised eye. Taxonomists (Standley, 1932) have so far failed to find
any essential morphological differences which would justify recogni-
tion of these forms as distinct. Chicleros, however, claim to know
the difference as soon as an incision is made in the bark. In the
writer’s experience there may be almost any degree of transition be-
tween the three forms, and frequently expert chicleros have been
very doubtful of the type when questioned about the exact identity
of certain trees.

In addition to the previously-mentioned forms of sapodilla, there
are numerous other laticiferous trees the latex of which is sometimes
used as chicle adulterants. Such adulterants, as far as is now known,
are derived chiefly from the Apocynaceae, Sapotaceae, Moraceae,
and Euphorbiaceae. The best chicle is reported to come from the
Mexican states of Quintana Roo, Campeche, and Yucatan, because
of their comparative freedom from these adulterants. In Peten,
Guatemala, other laticiferous trees occur in great abundance in the
chicle areas, and their latices have been used to dilute the increased
volume of good chicle. This is also true but to a less degree in Brit-
ish Honduras, when, during one rainy season, the writer collected
specimens of more than twenty trees, the latex of which is reported
to be used in varying degrees for adulterating good chicle. With the
view of bringing these datas together more concisely, the writer
has listed in tables 1 and 2 the species names, families, localities of
occurrence, and vernacular names of these adulterants. The order in
which they are arranged indicates the degree of frequency with

TABLE 1. SHOWING THE Sources, LOCALITIES, VERNACULAR NAMES, AND
LITERATURE REFERENCES OF CHICLE AND CHICLE ADULTERANTS IN
MEXICO AND CENTRAL AMERICA

Species

Achras zapota
( Chiquibul )

A. chicle

Calocarpum
mamosum

Calocarpum
viride

Dipholis Stev-
eEnsSONnIL

Dipholis sali-
cifolia

Bumela Guate-
malensis

Family

Sapotaceae

Sapotaceae
Sapotaceae

Sapotaceae

Localities and Vernacular
Names

British Honduras, Guatemala, Hon-
duras: sapote, chiqubull, chicle
bull, crown gum
Guatemala, British Honduras, Hon-
duras, Nicaragua, Panama: chicle
macho, sapote macho

West Indies: sapote, mamee sapote,
marmalade fruit (English). Mar-
tinique, Guadelupe: zsapotte,
grosse sapotte, sapote a creme
(French). Cuba: Mamey, mamee
sapote (Spanish). Mexico: tza-
potl (Nauhuatl), tspas Savani
(Zoque). Yucatan: zapote mamey
(Spanish), haas, chacal haaz
(Maya). Venezuela, Colombia,
Ecuador: Mamey colorado
(Spanish). Guatemala: Saltul
(Kekchi), tul-ul (Pokomchi),
Chul (Mame), chul-ul (Jacal-
teca). Costa Rica: bko (Cabé-
cara), kurok (Bribri), komkra
(Brunka), fm (Térraba). Pana-
ma: Oa-bo (Guaymi), mamey,
mamey de tierra. Philippine Is-
‘lands: chico-mamey (Spanish).

Guatemala: imgerto. Costa Rica:
sapote, zapote blanco (Spanish).
Honduras: gapotillo calenturiente
(Spanish). Salvador: sapote in-
gerto (Spanish). British Hon-
duras: red and white faisan. Nica-
ragua: sapote (Spanish). Hon-
duras: zapotillo (Spanish)

British Honduras: zapote faisdn
(Spanish)

Guatemala: dvalo, chaschin, acun,
chaxicaste

British Honduras: Mijico, Chachiga

Reference in
Literature

Pittier (1919) ;
Hummel (1925) ;
Record (1930) ;
Standley (1932)

Pittier (1914),
(1926) ; Pierre,
(1890), (1904) ;
Sloane (1725) ;
Jacquim (1760,
1763) ; Linnaeus
(1763) ; Miller
(1768) ; Gaertner
(1805, 1807) ;

Radlkofer (1882) ;

Cook (1913) ;
Standley (1925,
1928) ; Cook and
Collins (1903) ;
Popenoe (1920)

Pittier (1914) ;
Standley (1925,
1932)

Standley (1927,
1932)
Standley (1927)

Standley (1932)

Species

Bumelia lauri-
folia

Castilla fallax
and C. elas-
tica

Brosimum utile

Sideroxylon
amygdalinum

S. Gaumeri

S. Meyeri

Lucuma beliz-
ensis

Lucuma Dur-
landi

Lucuma sali-
ctfolia

Lucuma Hey-
deri
Lucuma cam-
pechiana
Stemmadena
Donnell-
Smith
Pseudolmedia
oxyphyllaria
Brosimum ali-
castrum

Family
Sapotaceae

Moraceae

Sapotaceae

Sapotaceae
Sapotaceae

Apocynaceae

Moraceae

Tasle 1. Continued

Localities and Vernacular
Names

British Honduras and Guatemala:
Silly Young (English), Suilion,
hoja largo

Guatemala: Ule, castiloa rubber,
castilloa. Mexico: arbol de bule,
hule, ule, olli, cwauchile, olcaguite,
ulcuagulil, ulcahwitl (Nahuatl).
British Honduras: hule macho,
tunu, toonu

Guatemala and Honduras: palo de
leche. Colombia and Nicaragua:
palo de vaca, palo de leche, cow
tree, arbol de leche, avichuri

Guatemala and British Honduras:
sapote faisan (Spanish), Sully
young (English)

Mexico: Caracolillo (Spanish).
British Honduras: Zoy, Dzo1i,
cream tree

British Honduras
zapotillo

Guatemala and British Honduras:
Silly Young (English), Suillion,
hoja largo, zapote (Spanish)

Guatemala: Zapotillo (Spanish)

and Mexico:

Mexico: zapote amarillo, zapote bar-
racho, zapote de nino (Spanish),
costiczapotl, atzapotl (Aztec and
Nauhuatl). Costa Rica: sapotillo
(Spanish). Guatemala : aceitunillo

British Honduras: Mamee cirulla
(Spanish)

British Honduras: Mamey cirera,
Mamey serilla

British Honduras: cojoton, cojon de
mico, cojon de caballo, chaclikin

Guatemala: wild cherry, mamba

British Honduras: Bread nut, mas-
ico, ramon (Spanish). Guatemala :
ramon, naranjillo. Mexico: ramon,
ojite (Spanish)

Reference im
Literature

Standley (1920)

Pittier (1909-1912) ;

Cook (1903)

Blake (1922) ; Pit-

tier (1918, 1926)

Standley (1925,
1929, 1932)

Standley (1925,
1932) ; Pittier
(1912)

Standley (1932)

Standley (1926)

Standley (1925,
1932)
Standley (1927)

Standley (1927,
1932)

Standley (1925,
1932)

Record (1930)

Record (1925,
1930) ; Pittier
(1918)

Species

Chrysophyllum
oliviforme

Tabernemon-

Tasle 1. Concluded

Localities and Vernacular
Names

British Honduras: Wild star apple
(English), chiceh (Maya), Chike.
Salvador and Honduras: Caimito
(Spanish). Salvador: zapotillo,
guayabillo (Spanish). Yucatan:
chiceh (Maya)

Apocynaceae British Honduras: cojon de pero,

Family

Sapotaceae

Reference im
Literature

Standley (1924,

1925, 1932) ;
Record (1930)

tana SP. cojoton
Thevetia Apocynaceae British Honduras: cojoton
nitida
Ficus lapathi- Moraceae Guatemala and British Honduras:
folia Kopo, mata palo, strangler fig
Ficus glabrata Moraceae British Honduras: wild fig. Guate- Record (1925) ;

Plumeria multi-

mala: higo. Salvador: Amate de
hijo grande
Apocynaceae British Honduras: sapilote

Standley (1917)

Record (1930)

flora

Cameraria Apocynaceae British Honduras: Chechem de ca- Record (1930)
belizensis ballo

Couma Apocynaceae Guatemala: palo de vaca; cow tree Record and Kuylen
Guatemalen- (1926) ; Karling
Sis (1935)

which, according to reports of chicleros in British Honduras, they
have been utilized in adulterating the good chicle from A. zapota.
This varies naturally in the different localities and countries ac-
cording to the occurrence of laticiferous plants, and there is, of
course, no universal agreement among chicleros and contractors
in this respect. The arrangement presented is accordingly personal
and tentative. Achras chicle gum and “chiquibul” are often collected
and sold, without mixing with A. zapota chicle, under the name of
Crown Gum in British Honduras. Contrary to the reports of many
chicleros, A. chicle or “chicle macho” in the writer’s experience
yields a goodly amount of latex in this colony, but its chicle is very
soit, difficult to mold, and resembles “‘chiquibul.’’ Chicleros, there-
fore, generally mix it with the gum of A. zapota in the proportion of
one to three, making a chicle that will mold and become quite firm.
The practice of adulteration in the jungle is not widely practiced at
present, since adulterated chicle can be readily recognized by tests.

TaBLeE 2. SHOWING THE Sources, LOCALITY, VERNACULAR NAMES, AND
LITERATURE REFERENCES OF CHICLE ADULTERANTS AND SUBSTITUTES
IN SouTH AMERICA AND THE Far East

Locality and V ernacular

Species Family Name
Couma utilis Apocynaceae Colombia: lirio
Manilkara sp. Sapotaceae Venezuela: pendare
(Mimusops)
Manilkara sp. Sapotaceae British Guiana
Dyera Low Apocynaceae Sarawak, British Borneo,

Sumatra, British Ma-
laya: dead Borneo, pon-
tianak, gutta jelutong

D. borneensis ©
D. Costulata ry
D. laxifolia i s

Alstonia Scho- i
laris

A. grandiflora

A. eximia

Rauwolfia * *
Spectabilis

Literature and
References

Vander Laan (1927)
Vander Laan (1927) ;
Pittier (1926)
Vander Laan (1927)
Vander Laan (1927) ;
Heyne (1914) ; Cor-
son (1927) ; Pearson
(1918)

Heyne (1914) ; Van-
der Laan (1927)

The price of such gum is accordingly reduced, and chicleros soon

discovered that adulteration is not profitable.

In these tables are included other laticiferous plants the gum of

which is used as substitutes, but which is nonetheless classed as
chicle in the countries where it is exported. The source of chicle is
less known in northern South America than in Mexico and Central
America, and much further study is necessary before definite state-
ments can be made with respect to the species of laticiferous plants.
According to Hoar (1924), over three million pounds of chicle were
imported from Colombia, Venezuela, Brazil, and British Guiana
annually immediately after the close of the last World War. This
quantity dropped considerably after political conditions improved in
Mexico and Central America, and according to later chicle import
data, it is considerably less.

According to Pell (1921), the largest amount and best chicle in
Colombia comes from the “zapote”’ tree, but whether this is 4. sapota
or some other member of the Sapotaceae is uncertain in view of the
wide range of trees which bear this vernacular name. Next in amount

and quality is “lirio” gum from various “‘lirio” trees, which is often
mixed with balata. The vernacular “‘lirio” is likewise extensively
used in Colombia and applied locally to many widely different plants.
Maloutia is a genus of laticiferous trees which occurs in the chicle
areas of Colombia, and Pell may possibly refer to a member of this
group. An anonymous writer (1921) and Vander Laan, however,
report Couma utilis, another species of the Apocynaceae which is
known locally as “‘lirio,”’ as the principal source of chicle in this coun-
try. Pittier (1918) describes Brosimum utile as one of the most
abundant sources of latex in Colombia, and it may possibly be used
as an adulterant. Species of Mamnilkara are also reported to be tapped
for chicle. These species are closely related to those which produce
the balata of commerce (Chevalier, 1932), and it is not improbable
that a considerable amount of latex from the latter, together with
that from species of Sapium, Sideroxylon, and Palaquim, is used in
adulteration. Along the north coast of Colombia is gathered an in-
ferior chicle known as “perillo,” which was exported to the extent
of nearly a half million pounds in 1923. Very little is known, how-
ever, of its source, as far as the writer is aware.

The chicle of Venezuela is known locally as “pendare” and was
exported to the amount of over a half million pounds in 1914, 1915,
and 1920. According to Fletcher (1927) and Vander Laan, it re-
sembles balata, and probably comes from a species of Manilkara.
Planchon (1888), however, reported that A. zapota is abundant in
the forests of Venezuela, but since his studies of the Sapotaceae were
made before the chicle industry had become extensively established,
he did not describe it as a source of gum. Doubtless, like Pittier
(1914), his description deals primarily with the cultivated sapodil-
las. Couma sapida (Pittier, 1926) occurs in the chicle areas of
Venezuela and may possibly be tapped for chicle.

Small amounts of chicle have been shipped from Panama, Costa
Rica, Nicaragua, and Honduras from time to time, but the exact
source of this gum is not certain from the literature. Doubtless, in
addition to A. sapota as a source, it comes largely from A. chicle and
the chiquibul form of A. zapota, and is adulterated with the latex of
other laticiferous species. In 1922 Costa Rica exported considerably
more than a hundred thousand pounds. The exports of Honduras
probably relate largely to Guatemalan chicle shipped through Hon-
duranian ports.

ECUADO
anor
40" EQUATO: i)
iD / I. C5
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Map 1. Areas of Mexico, Central America, and South America in which chicle
and chicle substitutes are reported to occur.

Very little is known concerning the source of Peruvian chicle.
Vander Laan reports it as coming from a species of the family
Apocynaceae. It is probably a mixture of various latices, since balata
and other gums occur in abundance in the same regions. Relatively
small amounts of chicle have been exported from Bolivia, Brazil,

0) 0OS/ oo Bt/ oo 9#/ ot t/ __oZ#/ __00#/ __oEY __o9E/ __ of E/ __oZE/_0OE/ _oB2/ __o92/,

‘poyoesj}xo St Suoynjaf yy ul eARTeWY puke SoIpUT sey IY} UI SeoIY “Zz AVIV

et) o22/_002/ __ol!__ 251) __obll__ off _20/! _080/ 290! oO! o20/ 000/86 095 _—otE

cSt! obb/ oct o0# BEL DEY hE = oCE/ OEY BCI oI oh oO BI oA ob oC OBO! «090! POS oO! (O0s—«i«iRG tC SCS

British and French Guiana, but the exact sources are not well
known. In Brazil occurs a species of the Apocynaceae which yields
the chicle known locally as Tamanqueira leiteira. Since Manilkara
and other laticiferous trees occur in great abundance here, this chicle
is undoubtedly a mixture. Part of the Brazilian exports probably
come from the eastern portion of Peru. In British Guiana a species
of Manilkara is reported to yield the chicle of commerce.

Another extensively used substitute is jelutong, which comes
from several species of the family Apocynaceae in Borneo, Sumatra,
and the Federated Malay States. According to Heyne (1914) and
Corson (1927), it is the product of various species, principally
Dyera Lowu, D. costulata, D. laxiflora, D. borneensis, Alstonia
scholaris, A. grandiflora, A. eximia, and Rauwolfia spectabilis.
Jelutong, which was formerly known under the names of “dead
Borneo,” “pontianak,”’ and “gutta jelutong,” is a soft pliable gum with
a resin content of seventy-five to eighty per cent and rubber varying
from nineteen to twenty-four per cent, according to Eaton and Den-
nett (1923). It is now being used extensively in the United States
for mixing with Achras zapota chicle, and according to Vander Laan
the total imports in 1910 reached fifty-two million pounds. Since that
time, however, it has dropped considerably, and in 1925 slightly
more than fifteen million pounds were imported.”

The various regions from which chicle, chicle adulterants, and
substitutes have been exported are shown in maps 1 and 2. These
maps have been made up chiefly from government and consular re-
ports and various articles on chicle, and with the exception of cer-
tain parts of Central America do not relate to actual observations in
the field by the writer. For this reason these maps will doubtless
prove inaccurate in many respects, particularly with reference to the
exact regions in which the latex-yielding trees occur, since the ports
from which chicle is exported are usually far removed from the
source.

(To be concluded)

2 Since the invasion of Malaya and the Dutch East Indies by Japan the
source of jelutong has been almost completely cut off.

More Fungi from the Front Lawn

Laura A. KoLtk

Since 1934, I have been recording the different species of fungi
which have appeared on the grounds of a small suburban home on
Long Island. Twenty-three species were reported in 1935 (TorRrEyA
35: 31-32, 1935), and since then the number has almost doubled ;
but it is as interesting to watch each year for the reappearance of |
the old “perennials” (?) as to welcome newcomers. Previous refer-
ence has been made to the two blue spruce trees, approximately
thirty years cld which dominate a portion of the front lawn. A scar-
let oak, a dogwood tree, and a hemlock, all about the same age as
the spruce, mark the boundary of the nearby adjoining property.
Other gymnosperms are scattered over the lawn, but they offer less
favorable cover for the growth of fungi.

Each year Russulas appear during the summer in the vicinity
of the oak. Short-stalked specimens are characteristic, so that the
purplish-red and grayish-green caps are in many cases scarcely
raised above the ground. Russula variata seems to be the common
species, and it is spreading in the grass beyond the immediate area
under the oak. Xylaria polymorpha, found in 1934, occurred in
abundance in the spring of 1936 in the form of small specimens
about an inch and a half tall, but seems to have disappeared. A maple
tree, growing too close to the oak, had been removed several years
previously and old roots may have been left in the ground, possibly
accounting for the appearance of the Xylaria in this location. How-
ever, this summer (1941) another specimen appeared, but in an
entirely different place. _

Amanitopsis vaginata var. plumbea is another species which
yearly makes its appearance in an area of approximately ten feet
between the scarlet oak and a narrow flower border along the side
porch of the house. The volva of this agaric sheathes the base of the
stipe much more closely than that of the heavier volva of Amani-
topsis volvata—a newcomer in the vicinity of the oak during the past
two years. The volva of the latter is very thick, and splits at the
margin into two or three deep clefts. The sporophore is quite slow
in reaching maturity, sometimes requiring two days or more to
emerge from the button stage. In July 1941, there appeared within
a few feet of the place where I have usually found A. volvata, an-

other agaric, with a large, bag-like volva (Fig. 1), deeply buried in
the soil, and of somewhat thinner texture. It was very similar to
A. volvata in its general characteristics, but was almost three times
as large as any specimens of that species which had appeared up to
date. The white pileus was covered with brownish scales, while the
margin was fringed with large loose flakes, similar to the covering

FicureE 1 (See Text)

of the six-inch stipe, which left a mealy deposit on the hands when
touched. Specimens of A. volvata showed a striate margin with no
indication of this fringe. Kauffman’s description’ of Amanita Pecki-
ana also fits this fungus in many particulars, but it will be necessary
to wait for its reappearance next year before a decision can be
reached. The specimen was not kept after photographing.

Laccaria amethystina reappears each year in the grass between
the oak and the dogwood tree, whereas the Amanitopsis species tend
to appear on nearby patches of bare soil. However, in 1940 another
Laccaria, L. ochropurpurea appeared on the bare soil, in the form
of a few depauperate specimens, but in September 1941, at least ten

1 Kauffman, C. H. The Agaricaceae of Michigan. Mich. Geol. & Biol. Surv.
Pub. 26, Biol. Ser. 5. vols. 1 & 2; 1918.

very well developed specimens appeared in a small area previously
occupied by the puff-ball Scleroderma aurantium. All these forms
are within a radius of 15 feet from the trunk of the oak tree. Typical
specimens of this Scleroderma have appeared each year, but several
smooth walled specimens were gathered the past summer which fit
Coker and Couch’s description’ of S. cepa, both as to peridium and
spore characters. A third species of Scleroderma has appeared in
sunnier situations in another area of the lawn. They are usually
much smaller than the specimens of the two already mentioned spe-
cies, and are replicas of those illustrated by Coker and Couch as S.
lycoperdoides in their Plate 94. The spores, however, correspond
more closely to those of S. tenerum on their Plate 120. I have col-
lected this species in the same areas of lawn since 1934.

The Boleti are represented by Boletus castaneus, which yearly
makes its appearance in the neighborhood of the oak, and by B.
chrysenteron, which has appeared from time to time in various
places on the lawn. A less frequent visitor is B. granulatus with its
stipe marked by reddish granular dots.

A species of Inocybe with angular, nodulose spores, has ap-
peared in 1940 and 1941 beneath a barberry hedge several feet from
the oak. Its cap, about an inch in diameter, shows the typical fibrous
markings of an Inocybe; it is umbonate with a dark umbo, and has
a tendency to split along the margin. This is the third species of
Inocybe to appear on the lawn. Two others, Inocybe infelix, and
Inocybe eutheloides (?) were found in 1934, but only J. infelix has
been a permanent resident. From spring to early fall this dingy
brown little agaric may be found on a barren patch of soil beneath
a rhododendron shrub. The spores of all three of these Inocybes
differ decidedly.

Ina patch of moss (narrow-leaved Catharinea) beneath the dog-
wood tree, a tiny yellow Clavaria has been found, and also Pleurotus
hypnophilus and a small white agaric possibly Omphalia gracillima
(?), but these are only among the occasional visitors.

Of the two blue spruce trees, which occupy a position directly
in front of the house, one seems to be much more favorably situated
for the growth of fungi than the other. Under the former, Amanita
muscaria has established itself permanently. Each year dozens of

2Coker, W. C. and J. N. Couch. The Gasteromycetes of the Eastern U. S.
and Canada. Univ. North Carolina Press. 1928.

specimens appear especially in the late summer and early fall. This
dangerous agaric is also found in other areas of the lawn, especially
under the hemlock and occasionally under a white pine. Of late these
Amanitas have produced caps which are more tan than orange in
color, but the volva is typical of A. muscaria.

This blue spruce harbors numerous other agarics beneath its
branches. The Inocybe with nodulose spores mentioned above, has
also been gathered here. Clitocybe infundibuliformis appeared in the
latter part of June 1936, and was found again in June 1937, and
May 1938. It has appeared since then, but no record has been kept.
The identification of a small gray Clitocybe has so far been doubtful.
Recorded as Clitocybe pinophila in 1934, other specimens gathered
since then, indicate it may be C. wilescens. A species of Psalliota
which appeared for the first time in the late summer of 1940, ap-
peared again in September 1941. I am inclined to think this is P.
abruptibulba. The fallen spruce needles, during a wet period in
1940, developed a conspicuous white mycelium which produced
brown sporophores two to three inches tall, with upward tapering
stipes, covered especially in the lower half with a dense white tomen-
tum, velvety to the touch, often binding several sporophores to-
gether at the base. This is Collybia (Marasmius, according to Pen-
nington®) confluens. The fetid Marasmius (MV. foetidus) appeared
twice on the lawn near the blue spruce; once in 1935, and again in
1937,

In the rear of the house, an apple tree occupies the center of the
yard, and for several years troops of Psilocybe foenescii were con-
tinually in evidence, but these have now disappeared. No other fungi
of interest have appeared in this area except a Psathyrella, single
specimens of which appeared several years in succession.

The following fungi have appeared only once: an Entoloma (spe-
cies undetermined), Naucoria semi-orbicularis, Hypholoma incer-
tum, and Lycoperdon Wrightu. Mutinus elegans (incorrectly re-
ported as M. caninus in 1935), Russula foetens, Coprinus micaceus,
Hypholoma sublateritium, Guepima (sp.) and a Lachnea-like As-
comycete (Patella albospadicea ?) all recorded in 1934, have not
reappeared. A Hebeloma was found recently near the place where
the so-called Pholiota aggericola appeared in 1934.

° Pennington, L. H. New York species of Marasmius, N. Y. State Museum
Bull. 179; Report of the State Botanist 1914.

The Zygomycetes are represented by Sporodinia grandis which
attacks the chestnut Boletus and also the Amanitas, and covers them
with a bright orange-yellow fuzz.

Even the rusts and smuts are represented in this limited area.
Several plants of smooth crab-grass in the back yard were infected
with Ustilago Rabenhorstiana and some Panicum dichotomiflorum
in one of the flower beds harbored the head smut Sorosporium Syn-
therismae. A tiny creeping Euphorbia had its leaves heavily rusted
with Uromyces proéminens in 1940. A special search for plant patho-
gens could no doubt have uncovered numerous others, since the
weeds above mentioned indicate neither a well-kept garden nor a
perfectly groomed lawn. For a mycologist, however, it is ideal.

BROOKLYN COLLEGE,
BrooKLyn, N. Y.

BOOK REVIEWS

About Ourselves

About Ourselves. By James G. Needham. The Jaques Cattell Press,
1941. Pp. XII + 276. $3.00.

It seems that a book so thoroughly publicized and by so popular
an author needs very little in the way of a review, especially when
it comes from a botanist with non-too-critical zoological leaning.
However, to those of us of the Torrey Botanical Club whose daily
task it is to present the biological aspects of human endeavors to
the young, a few words about the impressions made by this book
and the reasons why this book has such meaning for them, should
be of some interest.

The title “About Ourselves” may have many implications, but
since it has been written by a zoologist, one must naturally infer
that its discussions treat of the human being. Not only is this
true, but man’s relation to other animals and other human beings
are very much stressed. The book is accordingly divided into two
parts. The first deals with man in his biological aspect ; the second
deals with society in its biological aspects. The first part is replete
with topics which should appeal to the teacher in general and the
teacher of biology in particular. The language is not technical and

is adapted to the reading ability of an average intelligent layman.
Behavior, instinct, and learning are the subjects of some of the
most interesting chapters in Part 1. The second part is no less
interesting for such timely topics as the biological aspects of goy-
ernment, war, and religion are developed. The long list of readers
and commentators are high in praise of Needham’s efforts. Few
readers, however, have attempted to appraise the pedagogical value
of this volume. Briefly, as one teacher to another, let me say that
the author approaches his subject from the teacher’s point of view.
His story is told in a vein that makes it simple, interesting, and
often amusing. It is these characteristics that make many of the
topics models for teaching simple biological concepts to the non-
too-willing learner we meet in our schools today. The author
chooses from the known and non-technical subjects the facts best
suited to illustrate his point. The diagrams of Dr. Sargent are of
great simplicity and in two or three cases their purpose is not very
clear to the reviewer, except perhaps to heighten the basic nature
of the story.

The author at times seems to find it necessary to remind his
reader that “About Ourselves” deals with man in his zoological
aspects. For in such chapters as “Behavior,” and “Learning”’ little
is said about man. The chapter, “Nature and Nurture,” is interest-
ing and should serve as a review to all those who teach and find
little time for reading or experimenting. Here they will find a
slightly different outlook on the problems of heredity. Briefly
summarized in the author’s inimitable way when he tells the story
of germ plasm and body plasm, “Hats change but noses go on
forever.” In the chapter on the biological aspects of war there is
no outpouring of venom against the Axis powers but one finds
here an analysis of facts which lead to war and the contention that
war will be part of the untamed instincts and evil folkways of
Homo sapiens.

“About Ourselves” is a book which should interest a wide
group of readers, scientists, and especially teachers of biology as
well as those who are concerned with our present-day problems of

education.
MiIcHAEL LEVINE

LABORATORY DIVISION
MontTEFIORE HOSPITAL
NEw York City

The Microbe’s Challenge

The Microbe’s Challenge. By Frederick Eberson, Ph.D., M.D. 329
pages. The Jaques Cattell Press, 1941. $3.50.

To write a book on a scientific subject that will have appeal
for the lay mind as well as for the scientist is a difficult task and
yet this is seemingly what Dr. Eberson has achieved in his recent
publication, “The Microbe’s Challenge.”

Microbes are shown to have a way of living. They must grow,
eat, reproduce and die. The manner in which they set about this
business of living is vividly told. There are both good and bad
microbes and these are equally important to man. The microbial
parasite is a subject for contempt as are parasites in any walk of
life, but must be treated with respect because its parasitism is
necessary if it is to go on living. The fight to overcome these
disease-producing parasites is a fascinating one and puts to test all
of man’s ingenuity, as the author plainly shows.

The numerous disease-producing parasites or agents are each
described in detail and the means by which invasion is fought and
overcome clearly stated. One sees the body-producing poisons to
offset those produced by the microbe. Such words as toxin, anti-
toxin, bacteriophage, etc., are given meaning.

Virus diseases, the yellow fever problem, and many others
are set forth in a manner that will arouse enthusiasm for the
scientist and respect for the laboratory. The how and the why of
epidemics is but one of the problems met with.

Indeed the microbe’s challenge is being met with, and though
the path is hard and strewn with difficulties, much success has
been attained since Louis Pasteur first started out on the journey.

In addition to the above, the author gives a true and accurate
account of the history of the development of bacteriology and the

men who have made this possible. ROR Sete ae

Plant Anatomy

Practical Plant Anatomy. By Adriance S. Foster. D. Van Nostrand
Company, 1942. Pp. 155. $2.50.

If one approaches Dr. Foster’s new book as this reviewer did,
by way of the pre-publication announcement, the results are likely
to be disappointing. The publisher’s notice leads one to expect

another Eames and MacDamiels with all the recent findings in-
cluded and laboratory directions added. What one finds is a first
rate laboratory guide. The author defines his purpose as the
bridging of the gap between theory and practice in the study of
plant anatomy. The form of the book is admirably suited to this
purpose.

Each chapter consists of a discussion of the pertinent details
of modern theory concerning the topic considered and an outline
of practical laboratory exercises. ‘The first two chapters have to
do with the general characteristics of plant cells. The third chapter
is on meristems. Knowing Dr. Foster's excellent work in this
field one could wish that this chapter were more complete. The
various theories as to the structure of the apex certainly deserve
more discussion than they get here. A student doing the proposed
collateral reading at this point would easily be confused by the
various systems of tissue designation which he would encounter.
Chapter IV is a unique and very helpful presentation of the various
systems of cell and tissue classification. The charts relating the
origin, position, structural characteristics, and functions of different
cell types are perhaps the most valuable single feature of the book.
In chapters V through XI each of the principal cell types is con-
sidered in detail. In chapter XI there is a good discussion of the
distinction between sieve-tubes and sieve cells, the neglect of which
has led to confusion in some modern papers. The last three chap-
ters cover the stem, leaf, and root as tissue aggregates. There is
a very brief appendix detailing certain special laboratory procedures.

As a working laboratory outline this book should prove of great
value. The material is well organized and clearly presented. In-
structors will appreciate the designation of specific materials which
can be used for each exercise.

The principal criticism of this book is one which perhaps can
be equally well applied to the teaching of plant anatomy generally.
There is too great a tendency toward the purely descriptive aspects.
Anatomy is a justifiable study only in that it is a manifestation of
development either in the sense that the anatomy of an organism
is the ultimate expression of its morphogenetic pattern, or, what
is really the same thing, that it is a picture of the physiological
differentiation. As there is often a gap between theory and practice
in plant anatomy so does the descriptive approach make for a gap

between form and function. We should have preferred to see
what is really a developmental picture approached with a less static
outlook. However, this is merely a personal viewpoint. There is
much to be said for learning anatomy by this purer, more Spartan
approach. Any student who covers faithfully the material outlined
in this excellent book will certainly know plant anatomy, and know

it well. W. Gorpon WHALEY

BARNARD COLLEGE
CoLUMBIA UNIVERSITY

General Botany

Fundamentals of Plant Science. By M. Ellen O’Hanlon. F. S. Crofts
& Co., 1941. $4.25.

The numerous botanical textbooks of recent years are roughly
divisible into two groups: those that are the work of young sci-
entists and, like a spring freshet, have vigor, clarity of outline, and
force of presentation; then there are those other texts, the works
of botanists who have already won their spurs; and these, like a
mature stream, tap deeper reservoirs of knowledge and present
the subject set in its whole and proper environment. Happily,
“Fundamentals of Plant Science’ belongs in the latter group.

The book is divided into two parts. In traditional fashion the
first deals with such topics as “The Plant Cell,” “Leaves,” “The
Flower,’ “Fruits,” “Roots,” “Stems.” In the second half, after
a chapter on “Alternation of Generations,” the groups of the plant
kingdom are considered—the “Algae,” “Fungi and Their Allies,”
“Bryophyta,” “Pteridephyta,”’ etc. Following this, a thirty-five
page chapter is devoted to “Genetics’’; the next eighteen pages
deal with “Organic Evolution,’ and the final chapter takes up
“Botanical History.” This is followed by a glossary. There is
ample botanical nourishment between the covers of this volume for
the elementary student of general botany—very probably more
than he will assimilate in one year. This is true of all of our good -
texts and allows for discretion on the part of the instructor as well
as of the student. The scope and content are not markedly
different from those of other standard works.

Any new textbook of botany, following upon all of those already
published, should possess certain distinctive features. In this book

they are not hard to find. The author, whose work on the liver-
worts is well known, has presented an admirable brief account of
this group. The Anthocerotales are considered in a rank coordinate
with that of the liverworts and mosses and are taken up last in
the Bryophyta. In the Pteridophyta the Lycopods are discussed
first, where they really belong, and not last, as in most textbooks.
In a volume of slightly less than five hundred pages the author has
found opportunity to devote a page to apogamy and apospory,
another to the Gnetales, another to the embryo sac of the lily and
other atypical angiosperms. The chapter on genetics includes a
discussion of epistasis and of xenia. Other topics, such as tree
rings and their significance, artificial parthenocarpy, and hormones,
are not omitted.

Perhaps the most unusual characteristic of this book is to be
found in the references at the end of the chapters. The author has
had the courage to add to those time-honored and time-worn cita-
tions, so familiar in textbooks, selected new ones, many of them
readable articles in the current journals. The illustrations are
clear and well drawn and, mirabile dictu, practically all original.
Throughout the book, as well as in the glossary, the derivation of
botanical terms is given.

Conceivably a treatise could be written in clear diagrammatic
fashion, presenting facts and little more; but such a one would
hardly be worth its ink, like a picture without shading. Any worth-
while book reflects the personality of its author, and this is certainly
true of “Fundamentals of Plant Science.” The author projects
not merely her personality but also her philosophy into her writing.
There will be those who disagree with this philosophy, and who will
therefore prefer other texts. This volume is appropriately bound

in green.
2) Epwin B. Matzke

CoLUMBIA UNIVERSITY

Tudor Medicine
An Herbal [1525]. Edited by Sanford V. Larkey and Thomas Pyles.
Pp. xxiv + 86. 72 pp. facs. Scholars’ Facsimiles & Reprints, 1941. $3.50.
“Here begynnyth a newe mater the whiche sheweth and treateth
of ye vertues & proprytes of herbes the whiche is called an Herball.”
Just how new the matter was we cannot now say, the author not

having revealed himself. It is possible that it was the publisher’s
own compilation of current beliefs and was not due to the researches
of any scholar. In all events, this work, “imprynted by me
Rycharde Banckes . . . ye xxv. day of Marche. The yere of our
Lorde M. CCCCC. & xxv.”, seems to be the first book on herbs
printed in English. It was preceded, however, by Bartholomew
the Englishman’s De Proprietatibus Rerum, the seventeenth book
of which was devoted to plants and their uses; the English version
of this work was printed in 1495.

The medieval herbal was primarily medical. Descriptions of
the plants are secondary; in the Banckes herbal there are almost
none. It is a collection of information about the physiological
properties of plants, in 207 chapters arranged more or less alpha-
betically. There is intrinsic evidence that it is not the work of one
hand. Some plants are introduced twice, their names differently
spelled.

In their introduction to the present edition the editors damn it
by calling it “quaint, old-fashioned.” It is as quaint as our popular
medical works will seem 500 years hence and, naturally, as old-
fashioned. Perhaps this is only their way of saying that it is a
genuine product of the sixteenth century. Its main concern is with
the “aching of a man’s guts” and the “wicked winds” that trouble
them, and other parts and complaints not here mentionable. The
hearty (I had almost written lusty) freedom with which these
contemporaries of Henry VIII discussed such matters doubtless
accounts for the somewhat redundantly anatomical characteriza-
tion of the work (again by its editors) as “sinewy, muscular.” Ii
you wish to read, in modern print and spelling, how our remote
forefathers treated their intestinal and other troubles with prepara-
tions hot or cold to various “degrees,” moist or dry, laxative or
-“constipulative,” here is your opportunity. I refrain from further
quotation ; the book is easily available. There is little here of purely
botanical interest; and none of the imaginative power of vivid
description which illuminates old Bartholomew’s pages. Nor have
we any means of determining whether the work represents the best
medical science of its day. To judge from certain remarks of
Gerard some years later, many herbals of those times were com-
parable to our almanacs rather than to our textbooks.

The book was popular in its day, running into many editions
during the thirty years after its appearance. In spite of this, copies
are excessively rare. It is here presented both in facsimilie and
in “modernized” version, with an extensive introduction given
largely to bibliographic review. The printer has done a careful
job, the pages in facsimile are easily legible, and the small volume
is neat if not distinguished.

In the “modernized” version the spelling and punctuation are
changed to accord with modern usage, and explanatory notes are
added here and there. Although we are told that misprints in the
chapters have been corrected, we find “‘affodil,’ an obvious mis-
print for “asfodil”; the characters f and s being so similar. And
it is curious that “Abrotinum” is changed to “Abrotanum”’ in the
text, while other names (e.g., ““Aristologia’’) are not so treated.
The most puzzling feature introduced by the editors is the inclusion
in brackets after the chapter headings (which are transcribed
unchanged) of “corrected forms, as well as alternative forms which
might be of use to the modern reader in the identification of some
of the herbs.” I should have supposed that modern botanical names
would be of use here; but “Asfodillus” is changed to Asphodilus,
“Euforbium”’ to Euphorbium, and “Petrocilium” to Petrosilium ;
“Daucus creticus” is corrected (?) to criticus; and what are
Amarusca and Centumnodia? Surely the uninitiated reader is
entitled to an explanation of these scholarly mysteries. The work
concludes with an “Index of herbs and plants,” which is actually
an alphabetical list, without page numbers, of all the plant names
not used in the chapter headings. No indication is given of which

herbs are not plants. ih Wl Ree

New York BoTANICAL GARDEN
New York, N. Y.

IMIDSIEID) IesS) Oley Isls, CLs

Trip TO MisTarRE LAgBoraTories, MILiBurn, N. J.,
FEBRUARY 7, 1942

About twenty members and guests of the Club met at Mistaire
Laboratories, in spite of a rainy afternoon. They observed the unique
laboratory and greenhouse, where an attempt is made to control all
factors of plant growth while using natural daylight. Special exhibits
were arranged to explain the research and the methods used in rais-
ing ferns, orchids, and other plants.

All plants are started with exceedingly careful pure culture tech-
nique on nutrient agar of known composition within glass containers.
Among the exhibits in the planting room was a vibrator, used to
shake the seeds or spores being sterilized. Prothallia were trans-
planted from one tube to another with a platinum needle over a
flame. Flaming stoppers were snuffed out under a copper cone at-
tached to a standard. A’practical method for siphoning sterilized
solution from a large flask into hundreds of older tubes, in order to
adjust acidity and moisture, was shown. The air conditioning system
was so regulated that the pressure within the planting room was
greater than in the laboratory or outdoors. This kept unwanted germ
laden air from entering through cracks.

Growth and development of the plants is governed by automatic
controls of humidity, temperature, and light in the greenhouse. A
humidistat controls two humidifiers. Vaporized cold water is used
because it is more beneficial to living organisms than humidity
formed by heating. The temperature in the greenhouse is kept be-
tween 70° and 80° F. by means of an electric thermostat. If extremes
of 65° and 85° F. occur, an auxilliary thermostat causes a warning
bell to ring in the house. The temperature is also controlled by an
outer layer of Solex glass which eliminates most of the infra-red or
heat rays of the sun, and acts as an additional insulation in winter.
Summer heat thermostatically regulates a fine spray of water be-
tween the two layers of glass. Automatic recording instruments make
permanent graphs of temperature and humidity.

Within the greenhouse, nine photo-electric cells were function-
ing. One automatically controlled a large shade ; the second, the day-
light and green fluorescent lights; the third and fourth, the light
recording meter; others were for general study of light intensity.

The use of spectroscope to analyze wave lengths was shown. Row
upon row of tubes and flasks were seen on trays in the greenhouse.
The very young stages were in light positions of low intensity ; the
older plants were in brighter light, according to their maturity.

Visitors in the laboratory were shown working charts, graphs,
and records, as well as a filing system containing the histories of each
tube and flask. Three methods for determining the pH of solution
were of interest.

In the house, guests could observe under two microscopes and
several lenses, such materials as germinating orchid seeds, moss pro-
tonema, fern prothallia and young sporophytes, and nodules on
clover roots. Displays of living plants in culture tubes and flasks
were examined at leisure. The development of ferns was shown
from the spores and prothallia to sporophytes of different ages.
Great interest was shown in the proliferation of Polypodium aureum.
Among the native ferns were: walking fern (Camptosorus rhizo-
phyllus), climbing fern (Lygodium palmatum), purple cliff brake
(Pellaea atropurpurea), Hart’s tongue (Scolopendrium vulgare),
Dryopteris Goldiana, Dryopteris marginalis, and maidenhair spleen-
wort (Asplenium trichomanes). Another series showed the develop-
ment of various types of orchids. One round flask, containing a hy-
brid Billbergia or flowering pineapple, had been completely sealed
for six years.

In one room an original humidified bay window, enclosed in
glass, contained large ferns growing in deep soil and orchids in hang-
ing pots. An outdoor bird feeding shelf was built into the window ;
below set into recesses, were two aquaria. The window and aquaria
were heated by a concealed radiator, and were artistically lighted.

The pleasant afternoon was brought to a close with a television
program and refreshments.

MIsTAIRE LABORATORIES Ciara S. Hires

152 GLEN AVENUE
Mitizsurn, N. J.

‘po[vas Ayojorduros ‘savoc XIS 10} SUIMO.AS Uda

Svy YSVY [VU oy) ut ‘ojddvourd Surromoy 10 VIstod [le Prqay oy, ‘Spryo10 pur sutoy AYOYI—PIGIYXO OST 9uE ‘T DI]

PROCHEDENGSTOR Eh, CLUB

MINUTES OF THE MEETING OF FEBRUARY 3, 1942

The meeting was called to order at 8: 15 p.m. at the American
Museum of Natural History by the President of the Club, Dr. C.
Stuart Gager. Thirty-seven members and friends were present.

The minutes of the previous meeting were adopted as read.

The following were unanimously elected to annual membership :

Miss Marion Johnson, Rutgers University, New Brunswick, New Jersey;

Rev. Jos. Wittkoffski, M.M., Maryknoll, New York; Fr. Marie-Victorin,
Inst. Botanique, 4101 rue Sherbrooke, Montreal, Canada.

Mrs. R. B. Woodleton of 454 Seventh Street, Brooklyn, New
York, was transferred from Annual to Associate membership.

The President called upon the Chairman of the 75th Anniversary
Celebration Committee for a report. Dr. Karling reported that the
Committee had held three meetings. The celebration will be held the
week of June 22. It will open with a banquet on June 22 and will
be followed by scientific meetings during the week. Announcements
and invitations which had been sent to institutions have already
brought thirty-four responses. Seventeen delegates have been ap-
pointed.

The report of the Auditing Committee was made by Dr. Tre-
lease to the effect that the books of the Club had been examined and
found to be correct.

The Corresponding Secretary, Dr. Bold, reported on the per-
sonnel of the Standing Committees of the Club.

The President announced that a vacancy in the Council had been
created by the resignation of Dr. Chandler from her term ending in
1943. Dr. Chandler became a member of the Council upon her elec-
tion as second vice-president of the Club. Dr. Zimmerman was nomi-
nated for the position by Dr. Matzke. This was seconded by Dr.
Dodge and Dr. Zimmerman was unanimously elected to fill this
position on the Council.

The scientific program of the evening was presented by Dr. Nor-
wood C. Thornton who spoke on “The Mystery of the Potato Chip.”
The speaker’s abstract follows:

The potato chip industry had its beginning in the middle of the nineteenth

century. Today more than sixty million pounds of potato chips are sold annu-
ally requiring approximately four times this quantity of fresh potatoes to

provide for this demand. An industry requiring many tons of potatoes per day
demands almost unlimited storage facilities to prevent the potatoes from freez-
ing or even being chilled during the winter. One of the primary requirements
of potatoes to be used for potato chips is that the tubers contain a low amount
of reducing sugar. For it is this type of sugar, and not the total sugar content
of the potato, that is responsible for the color of the finished product. Artificial
“chips” have been produced by cooking filter paper soaked in dextrose.

As to be expected, reducing sugar accumulates at low temperatures, and at
5° C. we found this to occur quite readily in the twenty-five varieties tested.
However, the reducing sugar values of the potatoes stored at 7° C. were about
one-third of the values of those stored at 5° C. and the values of those stored
at 8.2° C. were about one-sixth of those stored at 5° C. Delaying the start of
cool storage after harvest retarded the rate of increase of reducing sugar at
5° C. so that after ninety days a lower sugar value was obtained than with
potatoes stored immediately after harvest. Also, storage temperature differing
only 1° C. caused differences in the rate of sugar accumulation in the potatoes.
Only potatoes of known history (i.e. the variety, the temperature of the soil at
harvest, time after harvest storage is started and temperatures held during each
period of storage, etc.) should be used in experimental work when attempting
to compare different varieties as to their suitability for potato chips.

The varieties of the Rural group were outstanding in maintaining low re-
ducing sugar values and providing chips of good color.

Following the discussion of the scientific program, Dr. Bold read
a communication from Dr. Small concerning tickets for the Sports-
men’s Show.

The meeting was adjourned at 9: 25 p.m.

Respectfully submitted,

Honor M. HoL_tinGHuRST,
Recording Secretary.

MINUTES OF THE MEETING OF FEBRUARY 18, 1942

The meeting of February 18 was called to order at 3.35 p.m.
in the Members’ Room of the New York Botanical Garden. Dr.
Chandler, the second vice-president, presided. Thirty members
and friends were present. The minutes of the preceding meeting
were accepted as read.

Mr. Arthur C. Riemer, Box 241, Delmar, N. Y., was unani-
mously elected to membership in the Club.

The resignations of the following were accepted with regret:

Gladys B. Goddard of 747 Dixie Lane, Plainfield, N. J.

Lora Bond of Wellesley, Mass.
Harley J. Scott of 3720 Avenue Q, Brooklyn, N. Y.

oy)

The scientific program of the afternoon consisted of an illus-
trated talk by Dr. Norma E. Pfeiffer on “Experiments in connec-
tion with Lily Breeding.” The speaker’s abstract follows:

Lilies, which show a great diversity in their interactions, are often self-
incompatible and seldom give natural crosses in the field. Seed set with
foreign pollen often gives rise to seedlings showing maternal characteristics
only. But pollen tubes from foreign pollen have been observed to grow less
rapidly than own pollen tubes.

Mechanical stimuli applied to the stigma of regal lilies in the absence of
pollen failed to induce seed setting, contrary to a popular idea. However,
the stimulus of pollen can be substituted for in capsule production by chemi-
cals, as shown by experiments on the Easter lily in 1937. Chemicals used
by different workers showed variable results in different species in inducing
formation of bulbs in the leaf axils. Naphthaleneacetic acid (Beale) gave
bulbs in a lingiflorum variety, but not in the Formosa lily, while colchicine
solutions (Emsweller) induced bulb formation in the Formosa lily, but not
in L. longiflorwm, the latter solution gives rise to polyploids. Formation of
aerial bulbs was induced accidentally by the speaker in L. longiflorum by
stoppage of growth of the main stem and through low temperatures.

Kodachrome slides were shown to illustrate a number of lilies with their
hybrids.

After the meeting adjourned at 4.30 p.m., tea was served
through the courtesy of the New York Botanical Garden.

Respectfully submitted,

Honor HoLtincHuRST
RECORDING SECRETARY

MINUTES OF THE MEETING OF Marcu 3, 1942

The meeting was called to order by the First Vice-President,
Dr. J. A. Small, at 8.15 p.m. at the American Museum of Natural
History. Thirty-eight members and friends were present.

The minutes of the preceding meeting were adopted as read.

The scientific program of the evening consisted of an illustrated
talk by Prof. Ralph Stewart on “Collecting Plants in Kashmir.”
The speaker’s abstract follows :

Kashmir is a very irregular bit of territory at the extreme north of India
where Afghanistan, Russia and Tibet come in contact with British India.
It is a native state, ruled by a Maharajah and extends roughly from 32 to
32 degrees north and 72 to 80 degrees east. It is all mountainous and very
rugged and except for the famous Vale of Kashmir is sparsely inhabited.

There are many elements in the flora because of the great altitudinal
range. Jumu is only 1,000 ft. above sea level and plants have been collected

100

up to 19,000 ft. north of the main range. In the foot-hill zone there is a
sub-tropical element with some plants which range as far as the Philippines.
In the Indus Valley there are desert plants ranging to the Mediterranean.
There are temperate and alpine plants which are found in various places as
far away as the Alps and others which apparently have come in from China.
Behind the Great Range the flora becomes like that of Tibet and Central
Asia.

Although the flora is most varied, the total number of species is probably
not more than 2,500 and in addition there are about one hundred ferns.

MaIn ZONES

I Sub-montane, largely varied types of thorny scrub.

II Pinus longifolia zone, 3-6,000 ft.

III Pinus excelsa and temperate hardwood zone, 6-8,000 ft.

IV Abies Webbiana zone, 8-11,000 it.

V_ Betula Bhojpattra zone, 11-12,000 ft.

VI Zone of shrubs, rhododendrons, willows, junipers, 12-13,000 ft.
VII Alpine meadows of herbs and grasses and sedges, 13-14,000 ft.
VIII High alpine zone of moraine and rock plants, 14-19,000 ft.

These zones apply to the Indian side of the main range of the Himalayas
and the altitudes vary a good deal according to exposure, rainfall, etc.

Behind the crest of the main range the forests disappear and closed forma-
tions are not common. Trees and crops have to be irrigated.

The meeting was adjourned at 9.35 p.m.
Respectfully submitted,

Honor M. HoLLtiInGHuURST
RECORDING SECRETARY

MINUTES oF THE MEETING OF Marcu 18, 1942

The meeting was called to order at 3.30 p.m. in the Members’
Room of the New York Botanical Garden by the Second Vice-
President, Dr. Chandler. Forty-seven members and friends were
present.

The minutes of the preceding meeting were accepted as read.

No changes in membership were reported.

The scientific program of the afternoon was presented by
Dr. Barbara McClintock who spoke on the “Contribution of the
Nucleolus to Genetic Investigations.” Dr. McClintock illustrated
her talk with slides and drawings.

101

The meeting was adjourned at 4.50 p.m. to enjoy the refresh- -
ments provided by the members of the Garden Staff.

Respectfully submitted,

Honor HoLLINGHURST
RECORDING SECRETARY

MINUTES OF THE MEETING OF ApRIL 7, 1942

The meeting was called to order at 8.20 p.m. by the President,
Dr. C. Stuart Gager, at the American Museum of Natural History.
Thirty members and friends were present.

The minutes of the preceding meeting were accepted as read.

The following were unanimously elected to annual membership :

Mrs. W. S. Randall, Alamo National Building, San Antonio, Tex.
Mr. Leon Tannenwald, 120 Lee Avenue, Yonkers, N. Y.

Mrs. Florence B. Cornish, Gillette, N. J.
Professor Hempstead, Castle University, New Haven, Conn.

The following were unanimously elected to Associate mem-
bership:
Miss Lilly Elkan, 39-89 46th Street, Long Island City, N. Y.

Miss Katherine L. Dudley, 509 West 122nd Street, New York City.
Mr. Zachariah Subarsky, 5450 Netherland Avenue, Riverdale, N. Y.

The resignation of Dr. James S. Wiant of 641 Washington
Street, New York City, from Associate membership was accepted
with regret.

Dr. Roger Wodehouse reported on the program planned by
the 75th Anniversary Celebration Committee. The celebration
will open on Monday, June 22, with a scientific program under
the chairmanship of Dr. Matzke at Columbia University. This
will be followed by a banquet at the Men’s Faculty Club. The
scientific program of June 23 will be held at the New York Botani-
cal Garden with Dr. Wm. Robbins acting as chairman. On Wed-
nesday, June 24, the program will be under the chairmanship of
Dr. Zimmerman at the Boyce Thompson Institute. Dr. C. Stuart
Gager will act as chairman of the meeting at the Brooklyn Botanical
Garden on Thursday, June 25. Field trips under the guidance of
Dr. Small will close the meetings on Friday and Saturday, June 26
and June 27.

102

Dr. Small announced that the Field Schedule would be in the
mails within a short time.

The scientific program of the evening was presented by Dr.
D. F. Jones of the Connecticut Agricultural Experiment Station.
Dr. Jones gave an illustrated talk on the “Chromosome Relocation
and Degeneration in Relation to Growth and Hybrid Vigor.” The
speaker's abstract follows:

The inherited characters studied by geneticists for the most part have
been highly selected to show clear-cut segregation with complete or nearly
complete dominance and recessiveness. Small changes in growth rate or
physiological efficiency are not easily detected and have been generally
overlooked but these play an important part in heterosis and give informa-
tion concerning growth and development. Spontaneous growth changes are
rarely found in maize endosperm but are easily identified and related to
chromosomal changes. Some of these give indication that critical regions
in the chromosomes are involved.

The meeting was adjourned at 9.45 p.m.
Respectfully submitted,

Honor M. HoLLtinGHuURST
RECORDING SECRETARY

103

NEWS NOTES

An extensive list of Institutions, Societies and Research Work-
ers in the pure and applied plant sciences in C. and S. America has
been prepared by the Editors of Chronica Botanica, in codperation
with the Div. of Agriculture of the Office of the Coordinator of
Inter-American Affairs, Washington, D. C. It has been published
in Chronica Botanica Vol. 7, No. 2 and 3 (March and May 1942).

Contributors to TorreyaA will please send all manuscripts to
Dr. H. W. Rickett, New York Botanical Garden, Fordham Station
P. O., New York, New York, until such time as a new editor of
TORREYA may be selected.

The current editor has been appointed chief drug specialist with
the Office of the Coordinator of Inter-American Affairs with Nel-
son Rockefeller and associated with the Board of Economic War-
fare. Fordham University has extended a.leave of absence for the
duration of the conflict and it is impossible to carry on the editorial
work associated with TorreEyA.

I want to express my appreciation to the officers of the Club
and to the many members who have cooperated in the publication
of TorrEYA.

WiLLiaAM J. BONISTEEL
EDITOR

THE TORREY BOTANICAL CLUB
OFFICERS FOR 1942

President: C. STUART GAGER

Vice-Presidents: JoHN A. SMALL, F.
CLyDE CHANDLER

Recording Secretary: Miss Honor M.
HoLLINGHURST

Corresponding Secretary: Haroxip C.
Bop

Treasurer: WW. GorpoN WHALEY
Editor: Harotp W. RIcKETT
Business Manager: MicHAEL LEVINE

Bibliographer: Mrs. LAzELLA SCHWAR-
TEN

Delegate to the Council, N. Y. Academy of Sciences: W. J. Ropsins

Representatives on the Council of the American Association for the
Advancement of Science:

A. E. HitrcHcock J. S. Kar_inc

Representative on the Board of Managers of the N. Y. Botanical Garden:

H. A. GLEASON
Council for 1942

Ex officio members

F. Clyde Chandler
Harold C. Bold

Honor M. Hollinghurst
W. Gordon Whaley

Elected Members

C. Stuart Gager
J. S. Karling
B. O. Dodge
John A. Small

1940-1942 1941-1942
Lela V. Barton Helen M. Trelease
R. W. Cheney R. C. Benedict
R. A. Harper J. H. Barnhart
E. W. Sinnott P. W. Zimmerman

Committees for 1942
ENDOWMENT COMMITTEE

Harold W. Rickett
Michael Levine
William J. Robbins

1942-1944
Edwin B. Matzke
Arthur H. Graves
J. M. Arthur
W. J. Bonisteel

Clarence Lewis, Chairman
Caroline C. Haynes
Henry de la Montagne

J. Ashton Allis
Helen M. Trelease

ProGRAM COMMITTEE

Harold C. Bold, Chairman (ex officio)
William J. Robbins
Edwin B. Matzke

Arthur H. Graves
W. Gordon Whaley
P. W. Zimmerman

Edward J. Alexander
G. G. Nearing
Vernon L, Frazee
Alfred Gunderson
Inez M. Haring

William J. Bonisteel
James Edwards
John M. Fogg, Jr.

Ferns and Fern Allies:
E. B. Bartram
A. W. Evans, E. B. Matzke
H.C. Bold
J. J. Copeland
A. H. Graves, J. S. Karling

J. W. Thomson, Jr.
R. Hagelstein

Mosses:
Liverworts:

Freshwater Algae:

Marine Algae:
Fungi:

Lichens:
Myxomycetes:

Fretp CoMMITTEE
John A. Small, Chairman

H. N. Moldenke
Rutherford Platt
Henry K. Svenson
Ellys B. Wodehouse
Eleanor Friend

Loca Frora COMMITTEE

Edwin B. Matzke, Chairman

Harold W. Rickett
Ora B. Smith
Herbert M. Denslow

Cryptogams

Robert Hagelstein
Michael Levine
Daniel Smiley, Jr.
Farida A. Wiley
James Murphy

Dolores Fay
H. Allan Gleason
Hester M. Rusk

R. C. Benedict, W. Herbert Dole, N. E. Pfeiffer

OTHER PUBLICATIONS

OF THE

TORREY aN cae CLU B

(1) BULLETIN

In addition to papers giving the results of research, each issue
contains the INDEX To AMERICAN BOTANICAL LITERATURE—a very
comprehensive bibliography of current publications in American
botany. Many workers find this an extremely valuable feature of the
BULLETIN.

(2) MEMOIRS

The Memoirs, established 1889, are published at pea in-
tervals. Volumes 1-18 are now completed. Volume 17, containing
Proceedings of the Semi-Centennial Anniversary of the Club, 490
pages, was issued in 1918, price $5.00.

Volume 18, no. 1, 108 pages, 1931, price $2.00. Volume 18, no.
2, 220 pages, 1932, price $4.00. Volume 18 complete, price $5.00.

Volume 19, no. 1, 92 pages, 1937, price $1.50. Volume 19, no.
2, 178 pages, 1938, price $2.00.

(3) INDEX TO AMERICAN BOTANICAL
LITERATURE

Reprinted monthly on cards, and furnished to subscribers at three
cents a card.
Correspondence relating to the above publications should be
addressed to
W. GorDON WHALEY,
Barnard College,
Columbia University,
New York, N. Y.

Ava £ Bedhlh | SO ae

Volume 42 July-August, 1942 Nia 4 NEW YOR

TORREYA

GARDED
A Bi-MonTHLY JOURNAL OF BOTANICAL NOTES AND NEwS

EDITED FOR
THE TORREY BOTANICAL CLUB

EAROED EH. CLuUM

John Torrey, 1796-1873

CONTENTS
Collecting Chicle in the American Tropics (Part 3)........ Joun S. Kariine 105
\

Book Reviews

Bible Plants for American Gardens.................. GerorGE T. Hastines 114

Work Book in General Botany..................... 00020000. W. H. Camp 115
Bicld Trips ofthe Clubs) 1 8 Oe ee RE Sa 118
INlewss Notesise tristan SS Me Bee ABU SOUL idee ceanrarn scan apt en euaiials 120

PUBLISHED FOR THE CLUB

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TORREYA

TorreyYA, the bi-monthly publication of the Torrey Botanical Club, was established
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W. H. CAMP DOROTHY J. LONGACRE

MEMBERSHIP IN THE TORREY BOTANICAL CLUB

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TORREYA

VoL. 42 JuLy-AvuGuUST No. 4

Collecting Chicle in the American Tropics
(Part 3)

JoHN S. KARLING

EvILs OF THE PRESENT METHODS OF COLLECTING CHICLE

It is quite obvious from the above description that the collection
of raw chicle is still in a very crude state. The sapodilla trees are
bled in a manner to secure the maximum yield at one time and at a
minimum of expense without much attention to conservation. No
extensive selection, planting, crossing, and grafting or budding of
high-yielding trees have been made, and no systematic production of
chicle on a large scale has been attempted. Practically all of the large
and important chicle areas in tropical America are controlled by the
governments and by large land-owning concerns and allotted to
chewing gum companies and individual contractors as concessions.
The general policy in the beginning was to grant only short-time
leases, and as a consequence the holders attempted to extract as
much chicle as possible while the concession was in their possession.
The granting of short time leases, the growth habits of the sapodilla
tree, and the nature of the countries in which it occurs have hindered
development of systematic chicle production in the virgin sapodilla
forests and the establishment of permanent centralized coagulating,
cooking, and supply camps. Furthermore, the fact that Achras
zapota can be profitably tapped only once within five to ten years has
contributed much to this condition of affairs and often made it un-
desirable and unprofitable for the small contractors at least to retain
their concessions for long periods of time.

In addition, the present basis of remuneration for tapping is a
great handicap to conservation and systematic production of chicle.
The chicleros are paid on the basis of the amount of gum extracted,
and as long as this system is in vogue, it is unlikely that they will
tap judiciously. However, experience has shown that this is the most
and perhaps the only practical basis of remuneration under the pres-

TorreEYA for July-August (Vol. 42, 105-120) was issued November 12, 1942,
105

106

ent jungle conditions and type of labor, since supervision in the more
or less inaccessible regions is almost impossible and would entail
considerable expense. As a result no particular premium is placed on
caution and care in tapping, and the only prerequisites of a chiclero
under the present system are the ability to climb with the aid of a
rope and make incisions which will secure the maximum yield of
latex and convey it without loss to the collecting bag at the base of
the tree. In their eagerness to secure the maximum amount of chicle
during the rainy season, chicleros often overtap the trees and cause
serious injury. Little regard is paid by the inexperienced chiclero to
depth of tapping and injury to the cambium. According to conserva-
tive chicle contractors, approximately fifteen per cent of the tapped
trees are eventually killed by the present native machete spiral
method of tapping. Hoar (1924) claims that twenty-five per cent are
killed. This estimate is based primarily on the number of dead trees
which may be seen in the chicle areas and does not, however, rep-
resent accurate annual counts and careful observation. To the casual
and inexperienced observer it would appear at first sight that the
number killed each year is appallingly high, since the chicle forests
contain a high number of dead standing trees, many of which bear
the tapping scars. This large number, however, represents the ac-
cumulation of many years, since the sapodilla tree, because of its
hardness, usually remains standing for several years after death and
decays very slowly. Furthermore, many trees die a natural death or
are killed by wood borers which enter after tapping. Consequently,
the number of dead trees in any particular chicle area is not an ac-
curate index of the number that is killed by tapping each year.
Nevertheless, the long machete used in the native system of
bleeding is a difficult tool to control with respect to depth of tapping,
and the cambium is often completely severed at the point of tangency
of the bole and the cut. Quite frequently the cambium is removed
with the chip of bark, and the wood is accordingly laid bare. Direct
exposure thus of the cambium and xylem to the tropical midday
heat often leads to a rapid drying out of the uninjured cambium and ~
cortex immediately adjacent to the exposed region. This drying out
may extend as much as an inch or more under the bark all around
the injury, forming a dead region many times as large as the original
exposed area. This is well illustrated by the trees shown in Figures
9 and 10. Immediately after tapping in October, 1927, the cuts on

Injury effects of the native system of tapping.

108

these trees were painted with white lead to prevent drying out as
much as possible. Figure 9 shows the condition in June of the fol-
lowing year. In the lower of the two cuts here shown the outer sur-
rounding bark has been removed, and the area of cambium and
xylem exposed at the time of tapping is indicated by the two streaks
of white paint. The size of this region as compared with surrounding
area is obviously several times smaller. Figure 10 shows a portion
of a smaller tree photographed a year after tapping when callus for-
mation had apparently just begun. Removal of the hard outer bark
showed a large triangular-shaped dead area of exposed wood. Injury
and exposure of the cambium obviously involves not only that por-
tion which is immediately injured at the time of tapping but in ad-
dition a considerable surrounding area. As a consequence, callus
formation must begin a considerable distance back from the border
of the original cut underneath the bark, as is shown in Figure 10.
If, on the other hand, the chiclero moderates the depth of tapping,
and the cambium is not exposed to drying, callus formation begins
very shortly in the incisions.

Another destructive result of the machete-spiral method of tap-
ping is that on the side of the tree where the oblique rows of cuts
intersect, a panel or zone is formed which is traversed by a zigzag
line or channel of cut bark (Figs. 2 and 3). Each oblique row makes
an acute angle where it intersects the one below and above, and as
a result this panel includes a large number of acute angles. If the
cuts are deep and injure and expose the cambium, and if subsequent
drying out at the angles is severe, the bark of the entire zone may
sometimes slough off, leaving bare an irregular panel running the
entire length of the bole, as is shown in Figure 11. Such exposed
areas require many years for healing, and in the meantime wood
borers and fungi may get in and destroy large regions of xylem and
cortex. Figure 12 shows a tree that has been killed by wood borers
subsequent to injurious tapping.

The ultimate death or recovery, rate of healing and bark re-
newal, however, are not dependent entirely on the depth and method
of tapping. The age, condition, and reaction of the tree itself play
a signficant role. Individual trees which have been carefully tapped
may show signs of severe injury and ultimately die, while others
which have been bled very severely may readily recover. This is well
illustrated in Figures 11 to 13. Although the tree shown in Figure 11

on Sarge sapodilla trees.

ing

Os
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ay
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c
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vo
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wn
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n
v
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=
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S
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I
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110

was severely injured, it is, nevertheless, recovering, as is shown by
the well developed callus on both sides of the injured panel. The
tree shown in Figure 12 was killed by the combined effects of tap-
ping and wood borers. Figure 13 illustrates a magnificent specimen
which has been tapped twice and shows no signs of permanent in-
jury. Shortly after this photograph was taken it was tapped a third
time and gave a large quantity of latex. In individual cases it is thus
difficult to predict the ultimate effect of tapping and the reaction of
the tree to injury.

There is little doubt, however, that the native machete-spiral
system of tapping is ruthless compared to the method employed on
Hevea braziliensis and is gradually killing a large number of trees.
This, together with the large amount of gum annually extracted and
exported from the chicle areas, and the apparent slow growth of
Achras zapota, is gradually exhausting the forests of “wild” virgin
sapodilla trees. Areas in which supply of chicle seemed almost in-
exhaustible a quarter of a century ago are thus becoming depleted.
On the other hand, there are many contractors who maintain that
the present demand and consumption is compensated by the rate of —
growth and healing of Achras zapota and that a sufficient number of
young trees come into profitable yield each year to offset to some
degree the long interval of time required for a tapped tree to heal.
The chief basis for their argument is that certain old chicle conces-
sions or areas have been yielding approximately the same amount of
gum for almost twenty-five years and that chicle exports have been
increasing steadily. To anyone familier with the conditions in such
areas and who has had intimate contact with the chicleros, it is ob-
vious that the task of maintaining the annual demand is becoming
more difficult each year. Chicleros must accordingly tap smaller and
younger trees each year to meet the demand, and it is not uncom-
mon to find trees as small as eleven inches in circumferences which
have been completely tapped. To the writer, who has spent several
years of observation and experimentation in tropical America, it is
obvious that the demand in normal times is greater than the annual
production of latex by the sapodilla trees in southern Mexico and
Central America, and that under present tapping methods and lack
of conservation a time will eventually be reached when the supply
is exhausted. Before this condition arrives, however, greater utiliza-
tion of favorable adulterants and chicle substitutes by chewing gum

111

manufacturers may doubtless establish an equilibrium between sup-
ply and demand of raw chicle and thus indefinitely postpone the
time of exhaustion.

Although it has been apparent for a number of years that the
present system is gradually depleting the chicle forests, no deter-
mined effort has yet been made towards conservation, and it is only
within recent years that serious attention has been directed to chicle
production on a plantation basis. Since the tropical forests of south-
ern Mexico and Central America contained at first seemingly in-.
exhaustible quantities of chicle that could be extracted at compara-
tively small expense, this attitude was to be expected, and it was
quite natural that no extensive effort was made to cultivate Achras
zapota. Sporadic small-scale attempts at cultivation have been re-
ported (Anonymous, 1923) from Mexico and the Far Fast, but
until two decades ago no significant efforts were made to cultivate
the sapodilla tree. The present status of the sapodilla tree as to meth-
ods of tapping, identification and selection of the best-yielding varie-
ties, plantation culture, etc., remind one very much of Hevea bra-
giliensis in the early years of the rubber industry. Years of rubber
gathering in the wild were necessary before the importance of plan-
tation production was realized, and then followed a long period of
experimentation with methods of tapping, propagation, and cultiva-
tion, with the result that the present highly specialized methods of
rubber culture finally emerged. In relation to plantation production
the chicle industry is at present in about the same stage as was the
rubber industry at the beginning of the nineteenth century, with the
important exception, however, that the sapodilla tree appears much
less, 1f at all, suitable for plantation culture than Hevea braziliensis.

BotTaANy DEPARTMENT
CoLUMBIA UNIVERSITY

Literature Cited

Anonymous. Chicle or sapodilla “gum.” Bull. Imp. Inst. 9:147-148. 1911.

Anonymous. Gathering chicle for the American gum chewers. Sci. Amer.
Suppl. 88:172. Figs. 1-3. 1919.

Anonymous. ‘Chicle cultivation in Mexico and the Far East. India Rubber
World 68:628. Three figs. 1923.

Bell, P. L. Chicle. Colombia. A commercial and industrial handbook. U. S.
Dept. Commerce. Sp. Agents Ser. 206:85-87. 1921.

Blake, S. F. Native names and uses of some plants of eastern Guatemala and
Honduras. Contr. U. S. Nat. Herb. 24:87-100. Pls. 29-33. 1922.

112

Brown, P. Civil and natural history of Jamaica. London, 1789.

Chevalier, A. Les vrais et les faux Balatas. Rev. Bot. Appliq. d’Agri. Trop.
12:261-282; 347-358. Pls. 8-11. 1932.

Cook, O. F. The culture of the Central American rubber tree. Bureau Plant
Industry Bull. 49:1-86. Pls. 1-18. 1903.

.A new generic name for the sapote. Jour. Washington Acad. Sci.

3:158-160. 1913.

. Nomenclature of the Sapote and Sapodilla. Contr. U. S. Nat. Herb.
16 :279-282. 1913.

, and G. N. Collins. Economic plants of Porto Rico. Contr. U. S. Nat.

Herb. 8:1-264. Pls. 13-60. Figs. 12-13. 1903.

Corson, T. Jelutong. Empire Forestry Jour. 6:47-55. 1927.

Coville, F. V. In W. E. Safford’s “The useful plants of the island of Guam.”
Contr. U. S. Nat. Herb. 9:369. 1905.

Dannerth, F. The industrial chemistry of chicle and chewing gum. Jour. Ind.
Eng. Chem. 9:679-682. 1917.

Durland, W. D. Chicle—source of chewing gum. Amer. For. & For. Life
33 228-229. 1927.

Eaton, B. J., and J. H. Dennett. Jelutong. Malayan Agri. Jour. 11:219-222.
1923.

Fletcher, S. J. In Vander Laan’s Production of gutta-percha, balata, chicle,
and allied gums. U. S. Dept. Commerce Trade Prom. Ser. 41. 1927.

Gaertner, C. F. Fructibus et seminibus plantarum. 3:129. Pl. 203. 1805.

Heyder, H. M. Sapodilla tapping in British Honduras. Emp. For. Jour. 9:107-
112. 2 pls. 1930.

Heyne, K. Wild rubber species in the Dutch East Indies. Rubber Recueil. 1914.

Hoar, H. M. Chicle and chewing gum. U. S. Dept. Commerce, Trade Inform.
Bull. 197. 1924.

Hummel, C. Report on the forests of British Honduras with suggestions for a
far-reaching forestry policy. London, 1925.

Jacquin, J. Enumeratio systematica plantarum. 1760.

. Selectarum Stirpium Americanarum. 1763.

Karling, J. S. Couma Guatemalensis as a possible source of chicle. Am. Jour.
Bot. 22:580-593. 1935.

Linnaeus, C. Species plantarum. Ist ed. 1753.

. Species plantarum. 2nd ed. 1762.

Melendez, L. E. La explotacion del arbol de perillo en Colombia. 1920.

Miller, P. The gardener’s dictionary. 8th ed. London, 1768.

Pearson, H. C. Crude rubber and compounding ingredients. New York, 1918.

Pierre, L., and I. Urban. Sapotaceae. Symb. Antill. 5:95-176. 1904.

Pittier, H. A preliminary treatment of the genus Castilla. Contr. U. S. Nat.
Herb. 13:247-279. Pls. 23-43. Figs. 1-54. 1909-1912.

. New or noteworthy plants irom Colombia and Central America 17.

Contr. U. S. Nat. Herb. 20:95-132. Pl. 7. Figs. 1-62. 1918.

. On the origin of chicle with descriptions of two new species of Achras.

Jour. Washington Acad. Sci. 9:431-438. 1919.

113

. Arboles y arbustos nuevos de Venezuela. Cuarta y quinta decadas. Bol.

Cient. y Tech. Mus. Com. Venezuela 1. Caracas. 1926.

. Plantas usuales de Venezuela. Caracas. 1926.

Pianchon, L. Etude sur les produits de la famille des Sapotés. Montpellier.
1888.

Plumier, C. Nova plantarum Americanarum. Paris, 1703.

Popenoe, W. Manual of tropical and subtropical fruits. New York, 1920.

Record, S. J. “Cow trees.” Trop. Woods no. 7:29. 1926.

. Local names of the woody plants of British Honduras. Trop. Woods

24:15-28. 1930.

, and H. Kuylen. Trees of the lower Rio Motagua Valley, Guatemala
Trop. Wood 7:10-29. 1926.

Resources of the Empire. Jelutong trade of British Malaya. Rubber, Tea and
Cacao Series. London. 1924.

. Jelutong industry in Borneo. Rubber, Tea and Cacao Series. London.
1924.

Sloane, H. A voyage to the islands of Madeira, Barbados, Nieves, St. Christo-
phers and Jamaica. 2:124-125. Pl. 218. 1725.

Standley, P. C. The Mexican and Central American species of Ficus. Contr.
U.S. Nat. Herb. 20:1-35. 1917.

. An enumeration of the Sapotaceae of Central America. Trop. Woods

4:1-11. 1925.

. New species of trees collected in Guatemala and British Honduras by

Samuel J. Record. Trop. Woods 7:4-8. 1926.

. Six new trees from British Honduras and Guatemala. Trop. Woods

11:18-22. 1927.

. Flora of the Panama Canal Zone. Contr. U. S. Nat. Herb. 27:1-416.

Pls. 1-66. Figs. 1-7. 1928.

. A second list of trees of Honduras. Trop. Woods 21: 9-41. 1930.

———. Additions to the Sapotaceae of Central America. Trop. Woods.
31:38-46. 1931.

Vander Laan, J. W. Production of gutta-percha, balata, chicle, and allied
gums. U. S. Dept. Commerce, Trade Prom. Ser. 41:1-72. 1927.

114

BOOK REVIEWS
Plants of the Bible

Bible Plants for American Gardens. By Eleanor A. King. The Macmillam
Co. 1941. Pp. 203. $2.00.

Probably everyone is familiar with some of the many Biblical
references to plants, from the first chapter of Genesis, where on the
third day of creation “the earth brought forth grass, and herbs yield-
ing seed after his kind, and the tree yielding fruit’’ down to New
Testament times when Jesus looking out over the mountain side
said: “Consider the lilies of the field, how they grow .. . even Solo-
mon in all his glory was not arrayed like one of these.”’ Probably
the majority of people reading of the lilies picture to themselves
Easter lilies, so it may come as a surprise to read that the lilies re-
ferred to were anemonies.

There have been numerous magazine articles on flowers and
trees of the Bible. The Journal of the New York Botanical Garden
in March, 1941, had an illustrated article on “Plants of the Holy
Scripture” by Miss King, and also a check list prepared in connec-
tion with the Garden's exhibit at the International Flower Show by
Dr. Moldenke of all plants mentioned in the Bible with the scientific
names of the plants as they are understood by modern students. But
no complete work of a popular nature on the subject has appeared
until this book by Miss King. In the front there is a paragraph of
appreciation of the help given by the staff of the New York Botanical
Garden, especially of that of Harold N. Moldenke. Comparing the
book with the scholarly study—Plants of the Bible—distributed in
mimeographed form by Dr. Moldenke, it is evident that this work
was drawn on to a large extent in the writing of the present volume.

As the title suggests the book is a gardeners’ manual with direc-
tions for growing the plants, outdoors or in, especially for those in-
terested in plantings or gardens for church grounds. But it is much
more, as it identifies the plants mentioned in the Bible, tells some-
thing of their characters, uses and meanings to ancient peoples.
Merely identifying the species is often difficult, not only because the
names used by the English translators were given by men un-
acquainted with the plants of Palestine, but also because the writers
of the Scriptures were not thinking in terms of botany, but used
vernacular Hebrew or Greek names that often referred to more than

115

one kind of flower. For example, the rose was some flower with a
bulbous root—tulip, narcissus, crocus of amaryllis—probably a gen-
eral term including all of these ; the lily of the field, as already men-
tioned, was an anemone (Anemone coronaria), but possibly included
all the wild flowers blooming on the hillsides ; apples were apricots,
quinces or oranges; the gourd that shaded Jonah may have been a
vine of the gourd family, though many students believe it to have been
the castor bean. Of course, for many of the plants named there is no
doubt as to the species—the Cedar of Lebanon, olive, fig, green bay,
palm, and some of the spices and plants used for perfumes or in _
making incense.

The book will be of great value to those who desire to devote a
part of their gardens to these plants of such sacred memories, to all
students of the Bible and to plant lovers generally. The dozen full
page plates illustrate a few of the plants and give some suggestions
for flower arrangements that combine beauty with religious sig-

nificance.
GeorceE T. HASTINGS

An Individual Botany Text

Work Book in General Botany. By H. C. Sampson. Harper & Brothers,
New York, 1941. 242 looseleaf pages. $1.75.

The subtitle of this publication is “A problem approach to plant
science through observation and discussion.” This, perhaps as well
as any single phrase, can be used to describe the method of instruc-
tion in the beginning course at Ohio State University under the im-
mediate supervision of Professor Sampson. It is inevitable that
many teachers of elementary botany may look with some disfavor on
this guide for it can scarcely be said to follow traditional lines. It is
therefore necessary that a little of its background be reviewed.

There has been much discussion concerning the method of in-
struction followed in that institution. In the first place, the beginning
student is not assigned a chapter in a book and told to return the
next day and “recite his lesson.” Also, there is no differentiation be-
tween lecture and laboratory sessions, for the students meet in the
same room with their instructor one hour a day, five days a week.
This provides the necessary continuity of topic and concept so sadly
lacking in many courses; it also establishes firm contact between

116

teacher and student. And it is this contact which permits freedom
of discussion.

It has been said, and with truth, that the method of instruction
at Ohio State is “a discussion in the presence of the material.” The
lesson, then, begins with a consideration of the problem. This is fol-
lowed by a study of pertinent material, interspersed with discussion
leading to primary conclusions and these, ultimately, to broader
biological generalizations. In recording his observations and con-
clusions, the student thus personally accumulates a basic textbook
of botany. It is obvious that many items, especially of a theoretical
or extended nature, or requiring too precise experimentation, cannot
be observed or discussed during the study period. For these the
orthodox text’ is assigned, as well as supplementary reading. In this
way the student is prepared for a further adventure into the general
subject of botany; at least he has been given some insight into the
methods of scientific reasoning based on experimental procedures.
Thus, by seeing, doing, recording and discussing, the student learns
the same facts he might otherwise memorize from a book. However,
at the same time he also acquires the habit of gathering and evaluat-
ing evidence, a mental trait which certainly cannot be cultivated by
the other method. |

There has been considerable argument that the use of a set of
drawings, complete except for the labels, does not cultivate the stu-
dent’s powers of observation. The writer of these notes is able to
take issue with this viewpoint for he instructed at Ohio State Uni-
versity during the decade of transition from the old to the new type
of instruction and watehed the method develop with considerable
interest, particularly as it influenced student reaction. In selected
classes having paired IQ ratings there was no decrease in effective
learning where prepared drawings were used. The advantage is that
they eliminate a lot of useless “busy work” which wastes time which
might more effectively be spent in examining the material or in dis-
cussion. However, the instructor should be cautioned that prepared —
drawings can never take the place of the actual material and that the
student must learn to study the material first, using the diagrams
or detailed drawings as a means of recording his observations.

* Textbook of Botany. By Transeau, Sampson and Tiffany. Harper &
Brothers. New York, 1940. ?

117

Therefore, regarding the use of unlabeled drawings such as are
an integral part of the Work Book, it should long ago have been
obvious that careful observation and accurate recording of scientific
information have but little in common with artistic ability. It has
been one of the major crimes of our biology teaching that we have
continued to penalize the student who is not congenitally an artist.
The argument that the professional botanist should be able to draw,
and therefore must learn in the beginning course, is certainly a fal-
lacy. Those who advocate this doctrine have somehow forgotten that
it is not the function of the introductory course to create professional
botanists but to teach botany. It is very doubtful if a group of stu-
dents—sounding for all the world like woodpeckers on a tin roof as
they vainly try to “stipple in the cytoplasm” with hard pencils—are
learning very much about the structure of protoplasm.

It would seem that I am defending Sampson’s Work Book. This
is unnecessary for it can stand on its own merits. But there are some
who further object to it on the ground that it contains too much ma-
terial, that they would not have time to cover all of it in a full year.
In general these are the same teachers who admit that they assign
a chapter in the text and then “hold the student responsible for every
word.” It is admitted that the Work Book does contain numerous
questions, but it should be obvious that it was not the intention of its
author and his collaborators that all of them be-answered. Certainly
many of them were introduced for the sole purpose of arousing dis-
cussion and to indicate the limits of our present knowledge, as well
as the need for more research before the question can adequately be
answered. It is perhaps a healthy mental attitude to instill in the be-
ginning student ; he should early realize that the science of plants is
not a closed subject and that much yet needs to be done.

There is also considerable complaint by some that the course does
not contain sufficient “morphology.’’ This unquestionably results
from the fact that the Work Book is not divided into sections labeled
“physiology” and “morphology.” There may be some lack of delving
into the more obscure of the “life histories” but the course actually
contains considerably more real morphology than is at first appar-
ent—probably more than most courses—for it is integrated with
the functional activity of plants, as it should be.

There is perhaps one drawback to a wider adoption of the course
as outlined in the Work Book. To teach it successfully, it is neces-

118

sary that the instructor have a fundamentally broad training in the
field of botany ; he cannot be a beginning graduate student interested
primarily in getting his degree, with his teaching a bothersome chore
to be sandwiched in at odd hours. He must know that the educator,
if he aspires to be worthy of the real meaning of the word, must do
more than stand in front of a group of students droning over phrases
which he has hastily snatched from a book a few minutes before class
time—and from the same text the students were supposed to have
“studied” the night before. The philosophical background of the
course has led to an organization designed to awaken in the student
an intelligent awareness of the nature of living organisms through a
study of plants. Under the guidance of a competent and sympathetic
instructor, this can be accomplished.

New York BoranicAaL GARDEN W. H. Camp
New York, N. Y.

IMUSILAD) WRIOES Ole Nas, CILIUIs
Trip oF NOVEMBER 2, 1941, ALONG THE APPALACHIAN TRAIL

Ten members and guests were present on this trip whose pur-
pose was to continue the botanical survey and census being made
by the Club of the New Jersey sections of the Appalachian Trail
maintained by the New York-New Jersey Trails Conference. In
the morning we covered the Dunfield Creek route from the Dela-
ware River to Sunfish Pond (Section la) and in the afternoon the
blazed route from Sunfish Pond back to the Delaware River (Sec-
tion 1), covering slightly over nine miles of trail in all. The
weather was intensely cold.

According to the official records in Dr. Small’s office there have
been identified thus far by Club members in Section 1 166 species
and varieties of spermatophytes, 11 pteridophytes, 4 bryophytes,
8 fungi, and 24 lichens. In Section la there have been found
159 species and varieties of spermatophytes, 17 pteridophytes,
15 bryophytes, 18 fungi, and 39 lichens. The total number of —
different species and varieties from both areas taken together is as
follows: spermatophytes, 238; pteridophytes, 19; bryophytes, 17;
fungi, 22; and lichens, 43.

Among the most interesting plants observed by us on our trip
through Section la were the American dittany (Cunila origa-

Wi)

noides), pubescent angelica (Angelica villosa), bearded short-husk
(Brachyelytrum erectum), eastern golden-saxifrage (Chrysosple-
mum americanum), beech-drops (Epifagus virginiana), large
coral-root (Corallorrhiza maculata), mockernut hickory (Carya
alba), bitternut hickory (C. cordiformis), and small-fruited hickory
(C. microcarpa), wild hydrangea (Hydrangea arborescens), downy
rattlesnake-plantain (Goodyera pubescens), ternate grape-fern
(Botrychium obliquum), cutleaf grape-fern (B. dissectum), com-
mon Virginia winterberry (Jlex verticillata), butternut (Juglans
cinerea), fringed milkwort (Polygala paucifolia), white swamp-
honeysuckle (Azalea viscosa), purple-flowering raspberry (Ruba-
cer odoratum), toothed whitetop aster (Sericocarpus asteroides),
vernal water-starwort (Callitriche palustris), common satin-grass
(Muhlenbergia mexicana), field basil (Clinopodium vulgare),
Torrey’s wild-liquorice (Galium lanceolatum), smooth rock-cress
(Arabis laevigata), hairy milkweed (Asclepias pulchra), deep-
green sedge (Carex tonsa), purple chokeberry (Aroma prunifolia),
and sheep-laurel (Kalmia angustifolia), all identified by foliar or
fruit characters, or, at least, in their post-anthesis stages. The
rare maidenhair spleenwort (Asplenium trichomanes) and walking-
fern (Camptosorus rhizophyllus) provided a thrill. Three species
were found still in bloom at this late date: the common bluets
(Houstonmia coerulea), American witch-hazel (Hamamelis virgini-
ana), and common white wood aster (Aster divaricatus). Large
quantities of a handsome earth-star (Astraeus hygrometricus)
were found along the trail and some mountain-laurel bushes were
seen to be infested with Phomopsis kalmuiae or Phyllosticta
kalmuicola.

At Sunfish Pond the most important finds were colonies of
the long sedge (Carex folliculata), dulichium (Dulichium arundi-
naceum), and sweet gale (Myrica gale). In Section 1, near the
Delaware River, we found fields filled by practically pure-stand
colonies of coralberry (Symphoricarpos orbiculatus), giving every
evidence of being native, some of the stands covering the major
portions of several acres. The European privet (Ligustrum vul-
gare), autumn oleaster (Elaeagnus umbellata), common tree-of-
heaven (Ailanthus altissima), Japanese honeysuckle (Nintooa
japonica), Japanese barberry (Berberis thunbergu), and European
barberry (B. vulgaris) were found as abundant escapes. Other

120

interesting plants observed were the green ash (Fraxinus penn-
sylvanica), maleberry (Arsenococcus ligustrinus), ebony spleen-
wort (Asplenium platyneuron), hooked crowfoot (Ranunculus
recurvatus ), northern wild-comfrey (Cyuoglossum boreale), moun-
tain-holly (Nemopanthus mucronata), common running-pine:
(Lycopodium clavatum), American trailing Christmas-green (L.
flabelliforme), early meadow-rue (Thalictrum dioicum), English
blue-grass (Poa compressa), pitch pine (Pinus rigida), common
wild-ginger (Asarum canadense), and American pennyroyal
(Hedeoma pulegioides). Particularly noteworthy were the soft
agrimony (Agrimonia mollis), white avens (Geum canadense),
low wild gooseberry (Grossularia hirtella), roughleaf bent-grass
(Agrostis hiemalis), smaller catspaw (Antennaria neodioica), and
plantainleaf catspaw (A. plantaginifolia). The liverwort, Cono-
cephalum conicum, was found in extensive mats on a moist cliff.
Along the river extensive beds of large-bracted plantain (Plantago

aristata) caused considerable comment. 4 d
H. N. MoLpENKE

NEWS NOTES

As announced in the last number of Torreya, Dr. William J.
Bonisteel is now chief drug specialist with the office of the Coordi-
nator of Inter-American Affairs. We regret that this has neces-
sitated his giving up the editorship of Torreya as he was well
qualified by temperament and experience to undertake such a task.
His work was well organized, and the issues of TORREYA were ap-
pearing regularly. He also had a number of ideas, which he had
not been able to put into practice, for improvement, and for mak-
ing ToRREYA more useful to the members.

The long delay since the last number of TorREyA appeared has
been due to the fact that there was no one immediately available to
take over the editorship in the absence of Dr. Bonisteel. Recently
Dr. Harold H. Clum has been asked to undertake this, and here-
after all contributions to TorrEyA should be addressed to him at
Hunter College, 695 Park Avenue, New York, N. Y.

Haroitp H. CLum
EDITOR

THE TORREY BOTANICAL CLUB
OFFICERS FOR 1942

President: C. STUART GAGER

Vice-Presidents: JoHN A. SMALL, F.
CLYDE CHANDLER

Recording Secretary: Miss Honor M.

Treasurer: WW. GorDoN WHALEY

Editor: Harotp W. RIcKETT

HoLLINGHURST Business Manager: MicHarEL LEVINE
Corresponding Secretary: Harotp C. Bibliographer: Mrs, LAzetta SCHWAR-
Boip TEN

Delegate to the Council, N. VY. Academy of Sciences: W. J. Ropsins

Representatives on the Council of the American Association for the
Advancement of Science:

A. E. HircHcock J. S. Kar_ine

Representative on the Board of Managers of the N. Y. Botanical Garden:

H. A. GLEASON
Council for 1942

Ex officio members

F. Clyde Chandler
Harold C. Bold

Honor M. Hollinghurst
W. Gordon Whaley

Elected Members

C. Stuart Gager
J. S. Karling

B. O. Dodge
John A. Small

1940-1942 1941-1943
Lela V. Barton Helen M. Trelease
R. W. Cheney R. C. Benedict
R. A. Harper J. H. Barnhart

E. W. Sinnott P. W. Zimmerman

Committees for 1942

EnDow MENT CoMMITTEE

Clarence Lewis, Chairman
Caroline C. Haynes
Henry de la Montagne

ProcGRamM COMMITTEE
Harold C. Bold, Chairman (ex officio)

William J. Robbins
Edwin B. Matzke

W. Gordon

FIELD COMMITTEE

John A. Small, Chairman
H. N. Moldenke
Rutherford Platt
Henry K. Svenson
Ellys B. Wodehouse
Eleanor Friend

Edward J. Alexander
G. G. Nearing
Vernon L. Frazee
Alfred Gunderson
Inez M. Haring

LocaL FLora COMMITTEE

Edwin B. Matzke, Chairman
Harold, W. Rickett

Ora B. Smith

Herbert M. Denslow

William J. Bonisteel
James Edwards
John M. Fogg, Jr.

Harold W. Rickett
Michael Levine
William J. Robbins

1942-1944

Edwin B. Matzke
Arthur H. Graves
J. M. Arthur

W. J. Bonisteel

J. Ashton Allis
Helen M. Trelease

Arthur H. Graves

Whaley

P. W. Zimmerman

Robert Hagelstein
Michael Levine
Daniel Smiley, Jr.
Farida A. Wiley
James Murphy

Dolores Fay
H. Allan Gleason
Hester M. Rusk

Cryptogams
Ferns and Fern Allies: R. C. Benedict, W. Herbert Dole, N. E. Pfeiffer
Mosses: E. B. Bartram
Liverworts: A. W. Evans,

E. B. Matzke
Freshwater Algae: H.C. Bold

Marine Algae: J. J. Copeland

Fungi: A. H. Graves, J. S. Karling, W. S. Thomas
Lichens: J. W. Thomson, Jr.

Myxomycetes: R. Hagelstein

PUBLICATIONS EXCHANGE COMMITTEE

Harold C. Bold, Chairman (ex officio) Amy L. Hepburn

Lazella Schwarten

OTHER PUBLICATIONS

OF THE

TORREY BOTANICAL CLUB

(1) BULLETIN

(2) MEMOIRS

The Memorrs, established 1889, are published at irregular in-
tervals. Volumes 1-18 are now completed. Volume 17, containing
Proceedings of the Semi-Centennial Anniversary of the Club, 490
pages, was issued in 1918, price $5.00.

Volume 18, no. 1, 108 pages, 1931, price $2.00. Volume 18, no.
2, 220 pages, 1932, price $4.00. Volume 18 complete, price $5.00.

Volume 19, no. 1, 92 pages, 1937, price $1.50. Volume 19, no.
2, 178 pages, 1938, price $2.00.

(3) INDEX TO AMERICAN BOTANICAL
LITERATURE

Reprinted monthly on cards, and furnished to subscribers at three
cents a card. 3
Correspondence relating to the above publications should be
addressed to
W. Gordon WHALEY,
Barnard College,
Columbia University,
New York, N. Y.

Ph os IF
Volume 42 September-October, 1942 Noes BO" |

NEW °

r A wD 2
NC

‘TVORREYA

A Br-MonTHLY JOURNAL OF BoTanicaL Notes AND NEws

EDITED FOR
THE TORREY BOTANICAL CLUB

HAROLD Fi, CLUM

John Torrey, 1796-1873

CONTENTS

Botanizing on Niue Island...................0.ceeceecceeecs T. G. YuncKer 121
Sedges and Rushes of Hot Springs National Park

aNd CNACINICY Aci) consis oe si haar Gin eto Ole ae re ee Francis J. Scutty 129
Cornus; Av Reply ass ae ee eae nee Outver A. FarRweLi 130
Gornus Asam es acs ee Meter hc nie ee nels SUI ee oo phare teres H. W. Rickett 131
Book Reviews

Standardized Plant Names...............20cceccecccecceeces A. B. Stout 132

ohn DOrre ys 6 ee es Ray ees e oe Coa ae LO: H. W. Ricxett 135

Methods of Plant Breeding......................... W. Gorpon WHALEY 137

Flowering Plants and Ferns of Arizona.............. CHARLES L. Gritty 138

An Introduction to the Study of Algae.................. Harotp C. Borp 140
BreldiErips of the Guba eee sey segs oe a ere a ha nasi ate Oa See orate 141
Proceedings’ of athe, Club). sia cick sscce sun rd otis sive ines aro o Unie a tag Mig ieee cies 145

ING yy s MINOLES Soe se rehe tse hed ete aac meer at HR Saal CRUE e keveee Sy claeel atattevevannne enue 151

PUBLISHED FOR THE CLUB

By THE FREE PRESS PRINTING COMPANY
187 CoLLEGE STREET, BURLINGTON, VERMONT

Entered as second class matter at the post office at Burlington, Vermont,
October 14, 1939, under the Act of March 3, 1879

TORREYA

TorreyA, the bi-monthly publication of the Torrey Botanical Club, was established
in 1901. TorreEYA was established as a means of publishing shorter papers and inter- ~
esting notes on the local flora range of the club. The proceedings of the club, book
reviews, field trips and news notes are published from time to time. The pages of
TORREYA are open to members of the club and others who may have short articles
for publication.

ToRREYA is iurnished to subscribers in the United States and Canada for one
dollar per year (January-December) ; single copies thirty cents. To subscribers
elsewhere, twenty-five cents extra, or the equivalent thereof. Postal or express
money orders, draits, and personal checks are accepted in payment. Subscriptions are
received only for full volumes.

Of the annual membership dues of the Torrey Botanical Club, $.50 is for a year’s
subscription to TORREYA.

INSTRUCTIONS TO CONTRIBUTORS

The manuscript should be prepared so that it conforms to the best practice as
illustrated by current numbers of TorrEYA. Manuscript should be typed double-
spaced on one side of standard paper. The editors may accept papers up to eight
printed pages in length. Longer papers may be published if the author agrees to bear
the cost of the additional pages. Illustrations (including tables and graphs) should
not exceed twelve per cent of the text; authors of more copiously illustrated ar-
ticles may be asked to pay for the excess material. Brief notes will be published
with especial promptness.

Drawings and photographs should be mounted on stiff cardboard and the desired
reductions plainly indicated. Figures should be so planned that after reduction they
will occupy the entire width of a page (4 inches) and any portion of the height
(6% inches). Labels should be parallel to the shorter dimension of the page. It is
best to combine illustrations into the smallest possible number of groups. Unmounted
material will not be accepted. Legends for figures should be typewritten and in-
cluded with the manuscript (sot affixed to the figures). All legends for one group
of figures should form a single paragraph. If magnifications are stated, they should
apply to the reduced figures.

TorrEYA is edited for the Torrey Botanical Club by

HAROLD H. CLUM
W. H. CAMP DOROTHY J. LONGACRE

MEMBERSHIP IN THE TORREY BOTANICAL CLUB

Manuscripts for publication, books and papers for review, reports of field
trips and miscellaneous news items should be addressed to:

' HAROLD H. CLUM
HUNTER CoLiece, 695 ParK AVENUE
New York, N. Y.

TORREYA

VoL. 42 SEPTEMBER-OCTOBER : No. 5

Botanizing on Niue Island

T. G. YUNCKER

To many, Niue Island has little significance because it is one
of the more isolated of the Polynesian islands, has had few Euro-
pean or American visitors, and about which there has been little
publicity. Furthermore, it lies in a lonely part of the south Pacific
ocean and has little or no strategic or commercial importance. Be-
cause of the manner of its geological formation and other features
it is, however, a very interesting island especially to the botanist
and the geologist. It is situated at 19° S. latitude and 169° 50’ W.
longitude with the Samoan, Tongan and Society groups as its
closest neighbors but with the nearest islands several hundred
miles distant. It has no harbor with safe anchorage during storms.
This fact discourages visits from any ships except the regular New
Zealand government service boat which, previous to the outbreak
of the war, was scheduled to visit the island once a month. For
weeks the only vessels one may see are the dugout canoes in which
the natives fish off the edge of the reef.

The island is approximately 13 miles long and 11 miles wide
and has a native population of about 4,000 which has remained
fairly constant for a number of years. The white population,
mostly government officials and their families, numbers less than a
score. The natives wear European clothing for the most part al-
though a wrap-around, skirt-like garment similar to the Samoan
lava-lava is also used. They are in the main a high type, quiet and
peaceful people. They do not, however, have the spontaneity of
the Samoans, for example, nor do they sing and play with as much
enthusiasm. This may be because life is considerably more difficult
in Niue than in most of the more fertile volcanic islands nearer the
equator.

Torreya for September-October (Vol. 42, 121 to 152) was issued January 29,
1943.

121

122

The first European to see the island appears to have been Cap-
tain John Cook who visited it on June 20, 1774. Cook and some of his
men, including Forster the botanist, attempted to land at two differ-
ent places along the western coast of the island, as is recorded in
his account of the visit. He was met with strong armed opposition
by the natives and consequently was compelled to retire without
exploring the island or collecting natural history specimens. Be-
cause of the violent resistance offered by the natives, Cook gave it
the name of Savage Island and by this name it appears on some
maps. The natives, however, do not like the name Cook gave their
island but prefer the native name of Niue which is said to have
been derived from niu, the Polynesian name for the coconut.

Most of the larger habitable islands of Polynesia are volcanic in
origin thus offering a diversity of elevation as well as other topo-
graphic features including streams, ravines, etc., commonly fa-
vorable for abundant plant and animal life. Niue, on the contrary,
is unique in that it is of the raised coral type of island. It has
been formed by the elevation of an original coral reef which was
nearly as large as the present island. The elevation was quite uni-
form so that the top of the island at present is nearly level or suff-
ciently so that differences can scarcely be noted without instru-
ments. There is a slight dip toward the center of the island sug-
gesting that the reef was originally of the atoll type. Following the
initial uplift a new reef varying from about one hundred to four
hundred or more meters in width developed around the raised part
of the island. Eventually the entire island experienced an addi-
tional elevation with the new reef now forming a shelf or terrace
about the original island. The total elevation of the island at
present is between 65 and 70 meters at its highest point. The edge
of the outer terrace ends at the sea with, for the most part, abrupt
and precipitous cliffs often 20 or more meters high. A new coral
reef, over which at high tide the waves dash against the rocky
cliffs, is now forming about the island. Perhaps the island has_
experienced other elevations but these two, at least, are apparent
to the geologically inexperienced eyes of a botanist. The geological
history of Niue should prove of great interest to students of the
raised-coral type of island.

During the centuries since the first elevation of the island, the
coral has been undergoing decomposition and change. Naturally,

123

the central or older part of the island, shows more disintegration
of the rock than does the terrace. With the coming of plant life,
humus has formed and soil has gradually accumulated, though soil,
in the ordinary sense, at no place exceeds much more than six inches
in depth. Beneath this top soil, which lies in pockets and crevices
of the rocks and is by no means a continuous layer, lies a layer
of decomposed coral limestone, white and powder¥, known as
makatea. This decomposed limestone will not alone support plant
life. It is now used to a considerable extent as a top dressing for roads
where, when rolled, it forms a compact and fairly permanent hard
surface. A red soil is also found in pockets here and there but to
a more limited extent. This soil when mixed with the top loam may
furnish a basis for plant growth.

The natives have cleared much of the island where the rock
has disintegrated sufficiently to permit working, but most of the
lower terrace, as well as large scattered areas on the upper level,
still remain too rough for the cultivation of crops. In clearing, the
felled trees and brush are burned which not only removes the
woody debris but also destroys some of the humus in the soil as —
well as reducing the microfloral content. The government offi-
cials are attempting to teach the natives not to burn their clear-
ings and to conserve all of the humus possible and that this is the
one great need of the soil. Seeds of Crotalaria anagyroides are
also supplied for scattering about the island in the hope that this
legume, which grows well under Niue conditions, may aid in soil
building.

Cultivation aside from simple hoeing is impossible. Taro,
bananas, yams, and sweet potatoes are planted in holes which are
made in the soil pockets with sharpened sticks. Weeds may be
pulled but for the most part the crops develop, if at all, without
benefit of mechanical aid. In spite of these handicaps, under nor-
mal moisture conditions, fair crops are produced. During periods
of drought, when the plantations fail, the natives are compelled
to subsist on coconuts, breadfruit, and the fruit, roots, and other
edible parts of wild plants of which there are not a great variety
providing usable parts. They may, of course, purchase a variety
of canned foods in the “bush”’ stores whenever they are able to se-
cure money through the shipment of produce to the New Zealand
markets.

124

The soil normally contains enough strength to permit only
about one season of cropping at a time. A new plantation area must
then be located and the old one allowed to revert to nature when
it soon becomes covered with a weedy second growth of herbaceous
and shrubby plants. After five or six years of such rest the soil may
again be cropped for a season. Thus, the soil is only able to produce
once out of every six or seven years. This represents a system of
rotating the soil rather than that of the crops. The difficulty of culti-
vating the plantations and the poor quality of the soil with the re-
sulting low crop yield obviously place a limitation upon the popu-
lation growth. Another feature which adds to the difficulty of liv-
ing on the island is the lack of any source of fresh water aside from
the rains. All water used for drinking purposes must be collected
from rains and impounded in reservoirs which, during times of
drought, may have to be severely rationed. Before the construc-
tion of the reservoirs the natives secured some water by catching
drippings from the roofs of caves. They also relied to a considerable
extent upon coconuts for drinking purposes.

As seen at a distance from the sea the island appears as though
completely forested with medium to large trees. This impression
is due to a considerable degree to the large numbers of coconut,
breadfruit, mango, and other introduced trees of economic use
which have been planted everywhere about the island particularly
in the vicinity of villages, all of which are situated on or near the
lower terrace.

A considerable proportion of the upper or older part of the
island has been cleared and cultivated at some time. Large areas,
however, still persist in what appears to be primitive forest. In
these areas the coral has resisted the forces of disintegration and
remains in exceedingly rough and rugged masses which make
walking off the trails exceedingly difficult. In places, deep, rugged
crevices and caverns with sharp projecting rock masses still exist.
These, now masked in some instances by lianas, ferns and other’

Figure 1. A forested “island” on the upper level with cleared plantation
area in the foreground in early stages of second growth. Ficure 2. A typical
native house in a grove of coconuts, breadfruit, bananas and other food and
ornamental plants. Ficure 3. The cliffs along the western shore near where
Captain Cook landed in 1774. The profile of the island in the distance shows
the upper level and surrounding secondary terrace.

126

growth offer some risk and persons have been injured or killed by
falling into them.

In these forested areas which have persisted because they are
too rocky to permit clearing and the cultivation of crops, one finds
a number of different species of shrubs and trees some with great
buttressed trunks and over a hundred feet in height. One also
finds several species of ferns and herbaceous plants and vines grow-
ing together in great profusion. Flagellaria gigantea, a curious
grass-like liana, which climbs to the tops of the tallest trees by means
of its prehensile-tipped leaves is also found here. Its great clusters
of small, white flowers are to be seen above the tops of the highest
trees and are collected with considerable difficulty. Among the
more abundant of the woody species to be found in the forested
areas and in the older thickets may be mentioned the following:
Celtis paniculata, Trema orientalis, Paratrophis anthropopha-
gorum, Ficus spp., Laportea Harveyi, Pipturus argenteus; Her-
nandia Moerenhoutiana, Pittosporum Brackenridgei, Adenanthera
pavonina, Inocarpus fagiferus, Micromelum minutum, Acronychia
sp., Canarium Harveyi, Dysoxylum Richi, Aglaia samoensis,
Glochidium ramiflorum, Macaranga Harveyana, Rhus taitensis,
Pometia pinnata, Ellatostachys falcata, Dodonaea viscosa, Colu-
brina asiatica, Alphitonia sizyphoides, Elacocarpus samoensis,
Grewia crenata, Psidium Guajava, Eugenia spp., Planchonella
samoensis, Diospyros, spp., Linociera pauciflora, Fagraea Ber-
teriana, Alyxia stellata, Tarenna sambucina, Psychotria insularum,
Morinda citrifolia, Morinda Forsteri, and Scaevola frutescens. The
cleared areas on the upper level are either bearing crops or are cov-
ered with a mass of second growth of ferns, grass, introduced
weeds, and low shrubs.

Along the coast on the exposed rocks or adjacent territory grow
a number of interesting native species. A conspicuous shrubby plant
which grows at the cliff edge is Bikkia grandiflora with its large
white tubular flowers up to six inches or more long. Large num-
bers of Messerschmidia argentea trees grow along the rocky cliffs
and when in flower are surrounded with great clouds of butterflies.
Among other common species found in this region may be cited:
Procris pedunculata, Pipturus argenteus, Hernandia ovigera, Cap-
paris sandwichiana, Leucaena glauca, Erythina variegata var.

127

orientalis, Mucuna gigantea, Aleurites moluccana, Triumfetta pro-
cumbens, Hibiscus tiliaceus, Thespesia populnea, Calophyllum
inophyllum, Pemphis acidula, Barringtonia asiatica, Terminalia
Catappa, Planchonella Grayana, Ochrosia parviflora, Cordia sub-
cordata, Heliotropium anomalum, Premna_ taitensis, Hedyotis
foetida, Gardemia taitensis, Guettarda speciosa, and Timonius poly-
gamus. Pandanus trees occur everywhere, the leaves of which are
much used by the natives for the weaving of large numbers of bas-
kets and other articles.

The government officials who have been stationed on the
island during the past forty years, together with the missionaries,
have been active in introducing ornamentals and plants of economic
worth. The natives themselves have also been responsible for the in-
troduction of a considerable number of species especially those of
more ancient introduction such as the coconut, banana, breadfruit,
yam, papaya, etc. In all of the villages are large numbers of those
ornamentals commonly found through the tropics including species
of: Crinum, Hymenocallis, Hedychium, Antigonon, Bougainvillea,
Bauhima, Clitoria, Acalypha, Euphorbia, Codiaeum, Hibiscus,
Polyscias, Plumbago, Jasminum, Allemanda, Plumeria, Nerium,
Cestrum, Thunbergia, etc. Salvia coccinea has become a common
weed about villages and along roadsides. In January it produces
a fine show with a blaze of red and pink flowers. Appropriately, it is
known locally as Bon Fire. Recently the Cassia shower trees have
been introduced and give promise of becoming an important dec-
orative addition. Occasional Norfolk Island pines and introduced
palms add to the landscape. The flame tree, Delonix regia, likewise
grows well and recently planted trees are flourishing and will soon
make a fine show of color during their blossoming season.

The writer was on the island for a number of weeks early in
1940 and attempted to secure as complete a collection of the plants
occurring there as possible. To that end I had the very hearty co-
operation of Captain William Bell, resident commissioner, and Mr.
Joseph McMahon-Box, then Secretary-Treasurer of the island
government, as well as a number of the natives. Good roads and
trails lead to all parts of the island so that nearly all regions were
readily accessible. In order to secure as many of the species as pos-
sible, particularly those which might be rare or obscure, the com-
missioner asked the native officials and older men of each village

128

to prepare a list of all of the plants which they knew. Many of the
older natives were known to have a wide knowledge of the plants,
particularly those with food or medicinal properties. The natives
gathered in groups in the various villages and each person con-
tributed the names of as many plants as he knew while one who
could write would list them. In this manner more than a dozen lists,
some of which included more than a thousand different native
names, were obtained. Naturally, there were innumerable duplica-
tions and it was necessary to match lists and combine names when
applied to the same species. From the list thus secured it was pos-
sible to seek those species not already collected and to locate, with
the aid of a competent guide, a number of species which would
likely have otherwise been overlooked. Finally, prizes were given
to natives who could bring plants which were rare and of which
specimens had not already been obtained. Most of those submitted
had already been collected but a few interesting additions were
secured in that manner. It was interesting to note that in many
instances the natives employed a workable form of binomial nomen-
clature to designate the various species, using descriptive specific
terms of color, habitat, leaf characters, etc. One wonders that
professional botanists waited until the time of Linnaeus before
adopting a similar system.

As might be expected, the endemic species are comparatively
few. Practically all of them occur also in Samoa, Tonga, Raratonga
or other islands of that general area. The total number of all species
collected was found to be less than 500 of which something less
than 50 percent are native. Considering the fact that there is little
variation in the thin soil, that the topography of the island is flat
and low, and that there is no surface water to furnish freshwater
habitats of stream or marsh, one should probably not expect a
larger number.

DEPauw UNIVERSITY
GREENCASTLE, INDIANA

WZ)

Sedges and Rushes of Hot Springs National Park and
Vicinity

Francis J. SCULLY

While making a collection of grasses of Hot Springs National
Park and vicinity a number of plants were collected which proved
to be the vegetative stage of many different species of sedges. Dur-
ing the next year a careful survey of this area was made for sedges,
attempting to collect them in the fruiting stage so the determina-
tions would be more accurate. The following forty-six species of
sedges represent this collection. Included also are twelve species
of rushes collected at the same time. The determinations of this
collection were made by E. C. Leonard of the Smithsonian Insti-
tution and E. J. Alexander of the New York Botanical Garden.

SEDGES
Carex blanda Dewey Carex triangularis Bock.
Carex Bushi Mackenzie Carex tribuloides Wahl.
Carex caroluuana Schw. Carex vulpinoidea Michx.
Carex cephalophora Muhl. Cyperus globulosus Aubl.
Carex comosa Boot Cyperus lancastriensis Porter
Carex crinata Lam. Cyperus ovularis (Michx.) Torr.
Carex debilis Michx. Cyperus pseudovegatus Stend.
*Carex festucacea Schkuhr. Cyperus refractus Engelm.
Carex Frankw Kunth. Cyperus rivularis Kunth.
Carex granularis Muhl. Cyperus rotundus L.
Carex Howeii Mackenzie Cyperus strigosus L.
Carex hystricma Muhl Cyperus Torreyii Britton
Carex mtumescens Rudge Eleocharis Engelmani Stend.
Carex laxiflora, Lam. Eleocharis lanceolata Fernald.
Carex Leavenworthi Dewey Eleocharis obtusa (Willd) Schultes
Carex lurida Wahl. Eleocharis tenuis (Willd) Schultes
Carex Meadu Dewey Fimbristylis autumnalis (L) R&S
Carex oxylepis Torr & Hook Fimbristylis puberula (Michx) Vail
Carex retroflexa Muhl. Kyllinga pumila Michx.
Carex rosea Schkuhr. Rynchosphora cymosa Ell.
Carex stipata Muhl. Rynchospora glomerata (1) Vahl.
Carex Swanu (Fernald) Mackenzie Scirpa lineatus Michx.
Carex tetanica Schkuhr. Schleria oligantha Michx.
RUSHES
Juncus acuminatus Michx. Juncus diffustmus Buchl.

Juncus aristulatus Michx. Juncus effusa L.

130

Juncus interior Weigand. Juncus tenuis Willd.

Juncus marginatus Rostk. Juncus validus Coville

Juncus setacens Rostk. Juncoides bulbosus (Wood) Small
Juncus scirpoides Lam. Juncoides campestre (L) Kuntze

904 MepicaL ARTS BUILDING
Hot Sprincs, ARKANSAS

Cornus, A Reply

OLiveR A. FARWELL

In Torreya Vol. 42:11-14 (1942) Dr. H. W. Rickett endeavors
to maintain as genera the subgenera Cynoxrylon and Eukrania pub-
lished as such by Rafinesque in Alsog. Am. (1838); the former
on p. 58 and the latter on p. 59. If Rafinesque were publishing
new genera, he would most certainly have made new combinations
or binomials under them. That the names were those of subdivi-
sions is proved by Rafinesque himself, who on p. 63 (1. c.) lists
and describes a species of Cornus as “281 Cornus (Eukrania)
cynanthes Raf. atl. j}. 151.” This can in no sense be construed as a
genus, Eukrania Raf. Aside from this we are not concerned with.
trying to interpret the ideas or unriddling the intentions of Rafin-
esque; but we are dealing with an actual fact in cold print. This
fact is that Rafinesque was monographing the genus Cornus and
creating new subdivisions thereunder; proved by the consecutive
numbering of the species under Cornus and not under the divisional
names. A perfect parallel is that of Chrysopsis of Nuttall under
Inula in his Genera I1 150, 151 (1818).

Many botanists, even Asa Gray, have considered Chrysopsis
of Nuttall as a well-published genus by him (1. c.), and have
credited Nuttall with the authorship of the binomials thereunder.
But it is no longer done as Nuttall listed his species under Inula.
Likewise as Rafinesque named his species under Cornus and not
under the new names, I have no doubt that botanists will treat them
as they treat Chrysopsis, as subdivisional names.

Box 265, LAKE LINDEN
MICHIGAN

131

Cornus Again

H. W. RIcKETT

Mr. Farwell’s contention really does not concern me, since I am
decidedly not endeavoring “to maintain as genera” the groups in
question ; this must be plain from my article. But Farwell’s words
prompted me to open yet again the Alsographia and thumb through
its pages, with results which the readers of this journal may find
as amusing as I did. For the first time I noticed on page 76 the
Index of Genera, the sub-title of which says “Subgenera in Italics.”
Ruaning my eye down this I quickly found Eukramia in italic and
Cornus in Roman, and was ready with chagrin to acknowledge my
error. But looking further I found Kraniopsis, which on page 58
is distinctly listed as “Subg.,’ in Roman, the same as Lentago,
while the sister subgenera Mesomera and Opulus are in italic.
Throughout the work the manner of listing the species is Rafin-
esque’s own. “Vib. L. rufidulum’ means Viburnum, subgenus
Lentago, V. rufidulum. On page 31 Rafinesque distinguishes
Leptaliv as a new genus to include the American species of
Fraxinus, and under it lists “Frax. vel. L. longifolia,’ and “Frax.
L. nuxta.” I think the conclusion is clear that one cannot solve
the riddle by typography, and I repeat that it is impossible to be
sure of the author’s intentions, especially since in several places
he very clearly implies that he had not completely made up his
mind on the status of these items. I am perfectly willing to refrain
from being dogmatic about the generic status of Eukrania or
Cynoxylon; but a weighty burden of proof must rest on anyone
who recognizes these genera but disregards these names. The
work, incidentally, is not a monograph of Cornus, but a supple-
ment to Rafinesque’s Trees and Shrubs of North America.

New York BotTanicaL GARDEN
New York, N. Y.

132

BOOK REVIEWS

Standardized Plant Names

Standardized Plant Names. Edited by Harlan P. Kelsey and William A.
Dayton, for American Joint Committee on Horticultural Nomenclature. Sec-
ond Edition. Pp. 675. Harrisburg, Pa.; J. Horace McFarland Company.
1942. $10.00.

The second edition of Standardized Plant Names is nearly twice
as large as the first edition, 1923. It was the aim to include in the
new edition the names of all plants of any economic or social value
to man and this has extended the total to “approximately 90,000
separate entries of plant and plant product names.” The new volume
is of primary value and interest in regard (1) to the standardiza-
tion of names and (2) to the “innovations” in the nomenclature,
the most important of which recognize the distinctions between (a)
true species (b) groups of hybrids (named “polybrids’). and (c)
clones.

The botanical names of genera and of their species are listed
alphabetically and the “approved” scientific names are printed in
bold-face type while synonyms or unapproved names are in italics.
Common, names for species and polybrids are in small captials as
are the names of clones. Names of polybrids are distinguished from
names of species by a symbol («) and the names of clones from
common names of species and polybrids by another symbol (¢),

In making decisions on approved scientific names there were
numerous collaborators and it is stated that it was the aim to apply
these names in accord with International Rules of Botanical
Nomenclature. In many cases when there is uncertainty in the ap-
plication of synonyms the authority is given; but authorities are not
cited for the names that are approved.

In any list of species names which is without either descriptions
or citation of authorities the identity of the group of plants to which
any name applies is not indicated. Hence the person who consults ~
Standardized Plant Names in regard to any name must either have
a knowledge of what that name applies to or be able to obtain this
information from other sources. If one has this knowledge for at
least one of the botanical names listed or for the one common name
that is given he can learn what the approved scientific name is.

133

For example, one learns that the generic name Amaryllis is pre-
ferred to the name Hippeastrum and that the species name Hemer-
ocallis Thunbergu is approved instead of the name Hemerocallis
serotina. In respect to the standardization of scientific names the
volume should be of value to gardeners and nurserymen.

In recognizing the clone and the polybrid the Editorial Com-
mittee of Standardized Plant Names renders a somewhat belated
service to both botany and horticulture. In the first edition these
distinctions were not made. That the rules of botanical nomenclature
adopted to date are inadequate in application to cultivated plants —
has been noted in various publications and also in the deliberations
and recommendations of the International Committee for Horticul-
tural Nomenclature.

It has long been recognized that all members of a clone have
collectively only the status of an individual. Methods of vegetative
propogation, especially for perennial plants, have made the clone
an important and very general horticultural unit. The term ‘‘clon”’
was proposed in 1903 but recently most writers have used the
spelling “clone.” The Editorial Committee of Standardized Plant
Names wishes to give the spelling that was first proposed prefer-
ence over that in recent general usage; but does not hesitate to
offer many new changes in the spelling and the pronunciation of
other terms.

The heterogenic nature of many groups of cultivated plants has
been emphasized by genetical studies as well as by the experiences
of gardeners. Often this condition arises after hybridization but it
is more or less developed in the population of any species. If seed-
reproduction is the rule for a group of hybrids, as in Petunia, there
is usually segregation into true-breeding varieties each of which de-
serves a distinctive name. But for most perennial plants the polybrid
group is soon separated into clones each of which deserves a clonal
name. In horticulture a polybrid group is a rather temporary and
variable group in comparison to the clone.

The horticultural varieties grown from seed are not listed in
Standardized Plant Names for certain genera; as, for example,
Petunia and Zinnia. But extensive lists of seed-grown varieties are
given for barley, oats, flax, rye, wheat, sorghum, and other agricul-
tural plants.

134

The Editorial Committee of Standardized Plant Names recom-
mends that there be “one standard common name for each plant.”
In reference to the names of species and true varieties the term
“plant” really refers to a group of individuals of successive seed
grown generations. When two or more common names are in use
for a group of plants only one is approved. Numerous new com-
mon names have been improvised. Numerous double names and hy-
phenated names in common use have been reduced to a single word ;
as, Lilyofthevalley, Jerusalemartichoke, etc.

There is much information concerning plants available in
Standardized Plant Names. For any genus of plants one may learn
how many species, varieties, polybrids, and clones are listed as 1m-
portant to man. In numerous genera the horticultural clones are
segregated and listed by common names and the names of the
originators are given (see Aster, Begonia, Azalea, Hemerocallis,
etc.). There is a list of plant patents with an index of the plants in-
volved. Lists are given of poisonous plants, range plants, state
flowers and trees, fiber plants, herb garden plants, and other groups
of plants that have special interest. These lists are useful as a basis
for obtaining specific information in descriptive literature.

In the designation of species, of clones, and of polybrids in
Standardized Plant Names there are numerous inaccuracies. Es-
pecially are many definitely recognized clones listed as polybrids or
even as species; but in most cases this treatment follows that of
some manual. This condition is illustrated in the nomenclature sug-
gested for the genus Populus. At this time this reviewer wishes to
record that the statement made in Standardized Plant Names that
he collaborated in deciding the nomenclature presented for the
genus Populus is an error.

Criticism of the volume is to some degree tempered when one
reads the following statements in the preface: “Standardized Plant
Names adopts the rule that species and natural varieties only are
entitled to Latin or botanical names and that all hybrids, clones,
polybrids, horticultural varieties and the like should receive suitable
English or common names. .... Time and other serious handicaps
make it impossible for the Editors to consistently carry out these
principles. Yet reasonable progress has been made and it 1s hoped

a later edition may see all necessary changes made in conformity
with this beginning.”

New York BoranicaL GARDEN A. B. Stout

The Years of John Torrey

John Torrey. A story of North American botany. By Andrew Denny
Rodgers, III. 352 pp. Princeton University Press. 1942. $3.75.

The journey of the Astorians during 1811 and 1812 began a ~
notable period in the exploration of western North America ;—_
notable for many reasons, among which we may reckon the pres-
ence of two well known naturalists. Subsequent expeditions (mostly
under the auspices of the United States Government) likewise in-
cluded natural history among the fields to be explored; the collec-
tion and description of the plants and animals and other products
of the country supplemented their purely geographical work. Speci-
mens flowed eastward in an increasing tide for identification and
preservation. Fortunately the prolixity of nature and the zeal of
collectors met their match in a few great naturalists who stayed at
home. Many North American plants went to William Jackson
Hooker at Kew; but the bulk of them during many years were
classified by John Torrey.

Torrey brought to this work acuity of perception and a talent
for organization (without which, indeed, it would not have been
brought to him). Though he was not himself a field botanist,
though he saw the western plants growing in their native places
only after his work was done, he labored to good purpose; his clas-
sification has formed an adequate skeleton on which to drape the
flesh of later research. His was a purely descriptive science. In-
quiries into the physiology of plants, into causes and first prin-
ciples, even into the Darwinian theories when they appeared, seem
to have interested him little. But in the scope of his knowledge, in
his mastery of detail, in his grasp of relationships, Torrey is en-
titled to first rank among the leaders in American botany.

Recognition was not slow in coming to such work, and both
labor and glory grew at the geometrical rate of the traditional snow-
ball. In his later years Torrey maintained a large correspondence
with botanists all over the world. He was instrumental in the es-
tablishment of the United States National Herbarium, and was one

136

of the first ““corporators” of the National Academy of Science.
From a group of young botanists inspired by his leadership grew the
Torrey Botanical Club.

From these remarks it is evident that a biography of John
Torrey must indeed be a “story of North American botany.’ Some
readers of the present work may feel, however, that title and sub-
title would better fit the contents if they were interchanged. Mr.
Rodgers has given us what is essentially a synopsis of the botanical
~ exploration of North America, with biographical details of the prin-
cipal American (and some foreign) botanists of the nineteenth
century ;—all against a background of extensive quotations from
Torrey’s letters. Some will think that a biographer should have
made a greater effort to penetrate this mass of detail and to portray
the human person within; others will doubtless maintain that the
letters tell the story. it 1s true nevertheless that the work is some-
thing of a hodge-podge, the main theme lost in the accompaniment.
This is the more to be regretted since, apart from his importance
to botany, Torrey was an engaging person; naive, religious, unsel-
fish, modest, shy,—and wholly lovable.

But there is another reason why this reviewer at least thinks
that the author should have written with a different emphasis.
Mr. Rodgers is not a botanist, and his attempts to evaluate the
place of Torrey in the history of botany are not to be taken seriously.
He assures us, for instance, that on two separate occasions Amer-
ican systematic botany was “born”; and it is rather astounding
to read that Mendel was one of the “great theorists, [who] built
on the vast taxonomic data gathered and organized by leaders such
as Torrey.” We see here a tendency evident in much modern biog-
raphy: to indulge in an orgy of hero-worship which covers a lack
of critical thinking. In the same tradition are the unfortunate at-
tempts at “fine writing.” As a substitute for creative literature we
are offered perfervid periods.

In spite of such shortcomings, the book has real value, par-
ticularly as a reference work for those interested in American sys-
tematic botany. The data are copious and accurate, and the student
will find useful notes on sources. There is also a “bibliography” of
Torrey’s works, from which dates of publication and other critical
bibliographical materials have unfortunately been omitted.

137

Style, after all, is a matter of taste; many will disagree with the
present reviewer in his strictures. But errors of grammar, punctua-
tion, and syntax are in a different category ; they are all too numer-
ous in this work, and contribute not a little to the peculiarity of
the style. One could wish that the editor of a University Press could
find time to attend to such small matters.

Tue New York BoraNnicaAL GARDEN H. W. RIcKeEtt

Plant Breeding

Methods of Plant Breeding. By H. K. Hayes and F. R. Immer. McGraw-
Hill. 1942. $4.00.

At a time like the present when it behooves every person to
examine his own endeavors and ask himself what he is contributing
to the nation’s war effort and to the cause of humanity this book
seems particularly pertinent and useful. It clearly serves the dou-
ble purpose of being a working guide for investigators in its own
field and an excellent review of the accomplishments and possibilities
of plant breeding for others.

Methods of Plant Breeding is a long book (well over four hun-
dred pages), but the subject of plant breeding is one of tremen-
dous consequence, and its accomplishments are already notable.
The first chapter is a brief statement of the role of plant breeding.
Chapters II and III cover respectively the genetic and cytogenetic
basis of breeding methods and the mode of reproduction in rela-
tion to plant breeding. The latter chapter includes a good practical
discussion of the heterosis question. In view of recent work of
Dobzhansky and others indicating the close association between
appearance and degree of hybrid vigor and the method of reproduc-
tion this arrangement seems particularly good. Chapter IV gives
details of methods for selfing and crossing the principal economic
crops. It is chapters like this one and later ones on the handling
of data which gives the book its value as a working handbook.
Chapters V, VI, and VII cover methods of breeding and Chapter
VIII correlates them with practical problems of breeding for dis-
ease and insect resistance. Chapter XIII returns to this discussion
of breeding for special characters. The intervening chapters are
given over to summary discussions of the genetics of wheat, oats,
barley, and flax. Chapters XIV and XV deal with breeding meth-

138

ods and the genetics of maize, which is genetically our best known
plant and probably the one in which breeding has so far obtained
the greatest improvements. Chapters XVI and XVII discuss con-
trolled pollination and seed production methods. The former in-
cludes a good section on the part played by incompatibilities and
sterilities in breeding problems. The last five chapters deal with the
standard methods of treating and analyzing data. A bibliography,
glossary of terms, and appendix of statistical tables complete the
book.

Methods of Plant Breeding could hardly have appeared at a
more opportune time. The plant breeder is to-day faced with what
is at once a challenge and a golden opportunity. Regardless of how
long or short the “duration” may be this country must for some
years to come produce both foodstuffs and other plant materials
to supply not only ourselves and our allies but later all those peoples
of the world whose lands have been devastated by war. This pro-
gram will necessitate further improvements in our main crop
plants, and the cultivation of many crops new to our agriculture.
The endeavor will be a tremendous one and this book should prove
a valuable guide to those entrusted with its breeding problems.

Finally it should be pointed out that several times the au-
thors emphasize that progress in the field, and its attendant bene-
fits to mankind, depend to a large extent upon free exchange of
ideas and materials among workers at different stations and in
different nations. This thought is one which it is to be hoped will
permeate fields far greater in scope than that of plant breeding.

BARNARD COLLEGE, W. Gorpon WHALEY
CoLUMBIA UNIVERSITY’

Apache-state Flora

Flowering Plants and Ferns of Arizona. By T. H. Kearney and R. H.
Peebles (and collaborators). Pp. 1,069, illustrated (29 plates and frontispiece)
and indexed. U.S. Dept. of Agriculture Misc. Publ. 423. May, 1942. $2.00. .

This, the second volume to appear in the last two years that can
be truly called a state Flora, takes its place alongside Deam’s Flora
of Indiana as an example for authors of future state Floras to emu-
late. The differences, other than format and general plan, between
these two state Floras are primarily due to the fact that while

12)

Kearney and Peebles have studied the Arizona plants, Deam has
lived with those of Indiana. This statement is in no sense a reflection
upon the Arizona authors and their comprehensive survey of their
state’s vegetation ; it is merely a summation of the differences in
“flavor” between the two volumes.

Certain of the families and genera of included plants, as in
Deam’s Flora, have been treated by recognized experts in these
groups; in this respect, as well as others, one may justifiably say
that the authors approached their problem in the “modern” sys-
tematic manner. Well written—that is as well written as any
manual, consisting primarily of keys, species-descriptions and
records of distribution, can be written—and rather adequately il-

- lustrated with definitely good photographs, the Flora also contains

an interesting discussion of the mantle of vegetation which, though
torn and frayed by climate and topography, covers Arizona. To one
who is addicted to maps as an aid to the interpretation of vegeta-
tional studies, a detailed map of the state, showing the major floris-
tic areas and accompanying F. Shreve’s discussion of vegetation
types, is a desideratum which might well have been included. An
outline map giving county limits, larger rivers and principal lo-
calities is a multiple guidepost to the “visitor” who dips into the
book.

Among the more interesting facts presented, at least to one
whose inclination is also toward things phytogeographic, is the pres-
ence in Arizona of two ferns, Asplenium exiguum and Cetararch
dalhousiae. The isolated Arizona stations listed, together with a few
localities for the former in northern Mexico, constitute the known
western hemisphere records of these two species whose primary
distribution is the Himalaya Mts., elsewhere in eastern Asia, and
Abyssinia.

As an indication of the scope and complexity of the flora of Ari-
zona, approximately 3,200 species, representing 128 families, are
treated, and the estimate is made that when the state is complete-
ly explored the total may well be more than 3,500 species. The
diversity of vegetation is due to several factors; among them the
altitudinal range and climatic variation, and the resultant com-
plexity of ecological habitats, within the state, as well as the num-
ber of primary vegetation-centers from which the components of
the Arizona flora have come.

140

All in all, then, the Flowering Plants and Ferns of Arizona is a
splendid contribution to North American botany. One can only re-
gret that in so few of these United States has the flora been so
thoroughly studied and so precisely depicted; it seems scarcely
necessary to say that the total complexities and coherence of the
vegetation of our country cannot be grasped so long as the distribu-
tion of a majority of its component elements, within so many of
the states, is adequately known. Cuaries L. Gritty

New York BoranicaL GARDEN,
New York, N. Y.

Algae for Undergraduate Students

An Introduction to the Study of Algae. By V. J. Chapman. Pp. 387.
The Macmillan Company. 1941. $3.75.

In the present volume the author has attempted to prepare a
short and relatively elementary text on phycology for undergraduate
students, hitherto available treatises being too unwieldly and com-
prehensive for such a purpose. The method of presentation, is in
general, the “type-method” in which one or more genera are se-
lected to illustrate the characters of each family. The book is
divided into fourteen chapters, including general chapters on clas-
sification ; reproduction, evolution and fossils ; physiology, symbiosis
and soil algae. Four chapters are devoted to ecology and distribu-
tion, and seven deal with the morphology of the type genera, fam-
ilies, orders and classes. References to important original sources
are included at the conclusion of each chapter. The logic of including
the Conjugales and Charales of the Chlorophyceae in the same
chapter with the Xanthophyceae, Bacillariophyceae, Chrysophy-
ceae, Cryptophyceae and Dinophyceae may be challenged in some
quarters.

Some curious inaccuracies pervade the book. For example: the.
plural of flagellum.is given as “flagellae” throughout the text. On
page 63, the Chaetophorales are referred to as a “family.’’ On page
72 it is implied that the oogonium of Coleochaete scutata possesses
a trichogyne. It is stated on page 102 that in Spirogyra “meiosis
takes place when the zygote germinates.” “Elachista” is written
for “Elachistea on page 145 ; the single egg of Desmarestia is referred
to as “ova” in figure 114. On page 30 species of Oedogonium with

141

antheridia and oogonia on different plants are spoken of as “dioeci-
ous homothallic’ while on page 21 Phacotus is described as a
“colourless unicell.”’ It is highly doubtful that any motile cells of
Botrydium have only one flagellum as they are figured in 83b. It is
regrettable that Juller’s (1937) important work on Stigeoclonium
is not referred to in the discussion of that genus, nor is it considered
in the general discussion of life cycles in the Chlorophyceae.

The last chapters on ecology and geographical distribution of
algae represent more or less of an innovation in phycological texts
in English, and the author is to be congratulated for having intro-
duced this material as well as a discussion of aspects of algal
physiology. Finally, the analysis of the derivation of the generic
names of the types described will be a helpful feature to many stu-

dents. Harotp C. Botp
BARNARD COLLEGE,
CoLUMBIA UNIVERSITY

IFVRIEID) WIRES Ole Walls, CLIVE

Trips oF APRIL 26 TO BUSHKILL FALis, PENNSYLVANIA

Thirteen members and guests of the Torrey Botanical Club
gathered at Bushkill Falls in the soft haze of an unusual morning
that in its warmth seemed like midsummer, but in its fragrance and
in the delicate green tracery of the new leaves it was definitely a
morning of early spring. Only the red maples in the low wet
grounds and the oaks on the drier hillsides faintly echoed the final
fanfare of the reds of autumn in the color of their expanding buds.

The group was honored this year by the presence and participa-
tion of Dr. Fulford, who contributed much to the study of the rich
Bryophyte flora of this area.

Many of the liverworts and mosses have been found and re-
corded on previous Torrey Club trips to this region (Torreya
40: 175-177; 41: 136-137). However, each year additional species
are collected, and a thorough search would undoubtedly yield very
many more. We had never identified Frullaria Asagrayana, with its
midrib-like ocelli, before; nor had we ever noticed the common
Chiloscyphus rivularis, which was growing in great abundance in
one of the small tributary streams. Not far away, also flourishing,

142

was Jubula pennsylvanica, coating the rocks of dark green. On the
sides of the main gorge, we had never seen the tiny Lejeunea patens,
the only slightly more conspicuous Leucolejeunea clypeata, nor
Jungermanma pumila. Directly opposite the main falls there were
miniature forests of Pellia sporophytes, their translucent stalks
glistening in the sunlight.

Antheridial receptacles of this year were well along on Mar-
chantia, but only archegoniophores of last year were in evidence,
somewhat the worse for wear.

Coptis trifolia, the Gold-thread, in flower, added its cheerful
touch to the dubious marshes, and the Fringed Milkwort was also
seen in bloom again.

Ceratodon purpureus, like other birds of passage, was roosting
in a burned over, waste spot.

On one of the drier hillsides, close to the path, Buxrbaumia
aphylla was growing more plentifully than we have ever seen
it, while in the brook below, the giant water moss, Fontinalis gi-
gantea, was still prospering, regardless of priorities.

The drive back through the village of Shawnee and the beauti-
ful valley of the Delaware led past apple and pear trees in full
flower and young grain fields in new green. It was still, on this
Sunday afternoon, a valley at peace in a world at war.

Epwin B. MAtTzKE

Trip oF JUNE 13, 1942, ro ENGLEWoop Cuirrs, N. J.

This Saturday afternoon trip covered a good botanizing region
only a half-hour’s bus ride from New York. Many of the common
plants of late spring and early summer, and numerous trees,
shrubs, and ferns were pointed out in relation to their varied
habitats of cliffs, open fields, woods, and swamps. Also some notes

were made concerning the geology of the region.
Mary Hoitzorr

Trip oF JUNE 12-13 To LAKE SHEHAWKEN, Pa.

This trip eventuated under several disturbing circumstances,
principally an unusually hot and humid Saturday, followed by a rainy
Sunday. The tour on Saturday took the party into Scott Township
about four miles north of Lake Shehawken. Among the more inter-

143

esting northern plants observed were Lycopodium annotinum, L.
tristachyum, Polystichum Braunu var. Purshu, Eriophorum calli-
thrix, and Cornus canadensis. A short side excursion was made to
see a field blue with blossoms of Scabiosa arvensis. Another walk
provided an infinite number and variety of Botrichium matricariae-
folium and B. lanceolatum var. angustisegmentum. Collections were
made of Polygonum natans var. Hartwrightw (not in flower),
Potentilla palustris and Salix lucida. The locality for Cetraria
islandica was also visited.

With the help of Mrs. Rodda of Palmerton, Pa., about sixty
species of birds were observed, among which were the Black-
throated Blue, Black-throated Green, Canadian, Blackburnian, and
Magnolia warblers, the Water Thrush, and the Veery and Hermit
thrushes. In one field was observed an unusual number of Henslow
sparrows, and one in particular which sat and sang (?) from the
roadside fence within a few feet of our car till we drove away and
left him still singing.

Besides the leader, the party comprised Mr. and Mrs. Rodda
and Mr. and Mrs. Hand of Philadelphia, Pa. A return visit under
a more favorable star is hoped for at a near opportunity.

W. L. Dix

Trip oF JUNE 20, 1942, Tro BRANcH Brook Park, NEwark, N. J.

Mr. Carl P. Witte, Horticulturist of the Essex County Park
Commission, accompanied the group through the Park telling the
people something about the plants of the Park and naming some
of the trees and shrubs for those interested. Dr. P. P. Pirone, Re-
search Specialist at the N. J. Agricultural Experiment Station,
pointed out a number of pathological conditions and gave us much
new information about the care and maintenance of shade trees.
Those participating were unanimous in declaring it an afternoon
spent to a pleasant and profitable advantage. Leader, Dr. Pirone.
Attendance, ten from Newark Museum Nature Club and Torrey
Botanical Club. ; Epwarp B. LANG

144

TRIP OF JUNE 20, 1942, TO THE FERN GARDEN OF Mr. AND Mrs.
W. HERBERT DOLE

The eighty-odd ferns and fern allies in this garden were tem-
porarily marked so that each species could be easily found and
identified. Most of the ferns in the garden have been growing here
from ten to twenty years and are well established and appear happy
in their present positions. A number are of recent introduction and
have been tried out for only a year or two. Several southern species
were planted only this spring and may prove unsuitable for this
latitude. The only way to settle that question seems to be to try
growing them.

My ferns all came through the winter in good condition, though
some species are always slow to start growth in the spring. Ferns
are more liable to damage by wind during the winter than by low
temperatures, and it has been found advantageous to protect those
in exposed locations with small branches anchored with pegs or
stones to prevent dislodgment. All are lightly covered with dry
leaves, except of course the larger local ferns which require little
attention.

The Cheilanthes ianosa on the limestone ledge in an exposed
position in full sun most of the day is still in fine condition and
shows considerable increase. From one small clump planted about
ten years ago there are now five clumps each larger than the original,
notwithstanding that a number of these ferns have been given to
other fern gardens. Woodsia ilvensis, also on the limestone in part
shade, after six or seven years is still in a very thrifty condition.
The Polystichums, set out in 1940 and given no special winter pro-
tection, are still doing well. These include P. anderson, P.
plumosum compactum, P. aculeatum plumosum (?) and P. vivi-
parum (said to be a West Indian fern). P. lonchitis, set out sev-
eral years ago, survived several winters then disappeared.

The Florida shield fern (Dryopteris ludoviciana) appears to be
hardy here. It has gone through three winters and shows increase
by offsets. Dryopteris celsa and D. chinensis set out a year ago are
growing nicely. The latter fern is especially attractive with its finely
cut lacy fronds. Several specimens of Scott’s spleenwort (Alabama
type) set out last year have developed new fronds and appear in
good condition. The same is true of Asplenium pinnatifidum planted

145

in a low wall of brown sandstone. Cystonuum falcatum, the holly
fern which I have mentioned several times in previous years, still
attracts attention with its shiny bright green fronds and exotic ap-
pearance. The alpine lady fern, Atherium alpestre var. americanum,
collected on Mt. Rainier and sent to me several years ago, appears
perfectly happy in its new habitat and has increased to several good
sized clumps. Blechnum spicant (Deer fern) has again developed
fertile fronds; last year there were only sterile fronds on this
northwestern fern. The small ““Mexican deer fern” which was sent
to me last year went through the winter without any protection
and is now larger than when it was received.

The afternoon provided ideal weather conditions and those who
came remained until late afternoon sitting in the shade to discuss
ferns and partake of refreshments provided by Mrs. Dole. At-
tendance 9. W. Hersert DoLe

ROCHA DIUNGS Ole Wells, CILiUI8

MINUTES OF THE MEETING oF Aprit 15, 1942

The meeting was called to order at 3.30 p.m. by the second
vice-president, Dr. Chandler, in the Members’ Room of the New
York Botanical Garden. Twenty-eight members and friends were
present.

The minutes of the preceding meeting were accepted as read.

The first portion of the scientific program was an illustrated
report by Mr. Libero Ajello on a New Chytrid Genus, Polychy-
trium. The Speaker’s abstract follows.

Polychytrium aggregatum is a new, polycentric, saprophytic species of
the family Cladochytriaceae which occurs in the decaying vegetation of bogs
in the ridges of Bearfort Mountain, Passaic County, New Jersey. It has a
coarse, richly branched rhizomycelium which becomes yellowish-brown at
maturity, and lacks spindle organs or intercalary enlargements. The sporangia
are smooth or tuberculate and produce spherical, posteriorly uniflagellate
zoospores which lack a conspicuous refractive globule but include a prominent
opaque lunate body. The sporangia dehisce by the deliquesence of the tip
of the exit tube or papilla. Dormant thick-walled resting spores have not
been observed, but the irregular tuberculate yellowish-brown sporangia are
strikingly similar to the resting spores of many Cladochytriaceous species.
However, they produce zoospores directly without going through a dormant
period.

146

The second talk was given by Dr. Edwin Matzke who spoke on
The Microscopic Anatomy in the Identification of the Commercial
White Pines. This was illustrated with slides and specimens. The
speaker’s abstract follows:

There are three common commercial species of white pine growing in the
the United States: the northern white pine, Pinus Strobus, the western or
Idaho white pine, P. monticola and the sugar pine, P. Lambertiana. In gen-
eral these trees are similar; the northern pine is distinguished from the others
by its finer needles, while the sugar pine can be told by its long cones.

The wood of these three species is also much alike in its gross as well as
in its microscopic characters. The texture is somewhat coarser in the sugar
pine, and the resin ducts are larger and darker in color. Sugar pine also has
the largest tracheids, northern white pine the smallest.

The most diagnostic microscopic difference between these three species
is the shape of the pits of the ray parenchyma cells. They are large and
oblong in P. Strobus, small, diagonally elongated and often apiculate or
lemon-shaped in P. Lambertiana, and intermediate between these two types
in P. monticola.

In many ways, microscopically as well as macroscopically, the western
white pine is intermediate between the other two. This is also true of its
distribution. However, other species also undoutbedly enter into this series.

The meeting was adjourned at 4.35 p.m. to be followed by a tea
served by friends at the Garden.

Respectfully submitted,

Honor M. HoLLtiINnGHURST
RECORDING SECRETARY

MINUTES oF THE MEETING oF May 5, 1942

The meeting was called to order at 8.20 p.m. by the second vice-
president, Dr. Chandler, at Schermerhorn Hall, Columbia Uni-
versity. Forty members and friends were present.

The minutes of the preceding meeting were accepted as read.

The following were elected unanimously to annual member-
ship:

Dr. V. E. Brown, Taylor University, Upland, Indiana

Dr. Wayne Manning, 14 Adare Place, Northampton, Mass.
Dr. Ernest Ball, Osborn Botanical Laboratory, New Haven, Conn.

The resignations of the following were accepted with regrev.
Dr. D. A. McLarty, Dartmouth College, Hanover, N. H.

Dr. George C. Wood, 4430 Tibbet Avenue, Riverdale, N. Y.
Dr. Walter T. Bedell, West Winding, Poughkeepsie, N. Y.

147

The scientific speaker of the evening was introduced by Dr.
Robbins. Dr. Beaman Douglas spoke on Botanizing in An Art
Museum and illustrated his talk with some very fine Kodachrome
slides.

The meeting was adjourned at 9.30 p.m. and was followed by
a tea served by members of the Columbia University Botany De-
partment.

Respectfully submitted,

Honor M. HoL_tiIncHURST
RECORDING SECRETARY

MINUTES OF THE MEETING oF May 20, 1942

The meeting was called to order at 3.30 p.m. in the Members’
Room of the New York Botanical Garden by the second vice-
president, Dr. Chandler. The minutes of the preceding meeting
were accepted as read.

The following were elected unanimously to annual membership:

William E. H. Schneider, Jr., 90 Engle Street, Englewood, N. J.

Prof. Seville Flowers, University of Utah, Salt Lake City, Utah
T. Monroe Kildow, Box 520, Tiffin, Ohio

The following was unanimously elected associate member :
Eleanor Ruth Witkus, 61-19 Grand Avenue, Maspeth, N. Y.

The resignations of the following were accepted with regret:

Don. E. Eyles, Memphis, Tenn.
Clifford S. Leonard, 31 Cliff Street, Burlington, Vt.

In response to the question raised regarding the progress of the
committee on the per capita cost of membership in the Club, Dr.
Dodge stated that he was awaiting a report from the Treasurer.

Dr. Bold moved that the Treasurer be instructed to pay for the
‘75th Anniversary Celebration Banquet dinners of the officially ap-
pointed delegates and speakers from outside the metropolitan area.
This was seconded by Dr. Karling and passed by the Club.

The chairman of the 75th Anniversary Committee, Dr. Karling,
announced that 97 institutions had appointed delegates to the meet-
ings.

The scientific speaker of the afternoon was Dr. W. H. Camp
who spoke on “The Genetic Structure of Populations and the
Delimitation of Species.”

148

Following the discussion of the talk, tea was served by friends
at the Garden.
Respectfully submitted,

Honor M. HoLtLtinGHuRST
RECORDING SECRETARY

MINUTES OF THE MEETING OF OCTOBER 6, 1942

The meeting was called to order at 8.40 p.m. at the Brooklyn
Botanic Garden by the President, Dr. C. Stuart Gager. Thirty
friends and members were present. The minutes of the preceding
meeting were accepted as read. |

The following was unanimously elected a sustaining member:

Thomas C. Desmond, 94 Broadway, Newburgh, N. Y.

The following were elected unanimously to annual membership :

Gladys Boughton, 448 Washington Street, Brooklyn, N. Y.
Margaret S. Rogers, 20 Haslet Avenue, Princeton, N. J.

Dr. F. L. Wynd, University of Illinois, Urbana, Ill.

Dr. Arnold Rocha, Rua Angelo Agostino, 18, Rio de Janeiro, Brazil
Dr. Selman A. Waksman, N. J. Agric. Exper. Sta., New Brunswick, N. J.
Dr. Joseph Austin Miller, 364 Prospect Street, South Orange, N. J.
Dr. John N. Martin, 507 Welch Avenue, Ames, Iowa

Nettie M. Sadler, 503 Allen Street, Syracuse, N. Y.

Dr. William A. Beck, University of Dayton, Dayton, Ohio
Clarence R. Hanes, Schoolcraft, Mich.

Dr. Thomas S. Stewart, 18th and Rittenhouse Square, Phila., Ba,
Francoise A. Kelz, 31 Dobbs Terrace, Scarsdale, N. Y.

Joseph Ravizza, 312 Stanley Street, New Britain, Conn.

Arthur M. Scott, 7035 Chestnut Street, New Orleans, La.

Rey. F. J. Mahoney, S.J., Regis College, Denver, Col.

Mrs. George H. Sinden, Vassar College, Poughkeepsie, N. Y.

Transfer from annual to associate membership was approved
for:
Dr. Alexander V. Tolstoouhov, 24 Arden Street, New York City

The following resignations were accepted with regret:

Fred. A. Barkley, Montana State University, Missoula, Mont.
Helen Berdan, London, Ont.

Mrs. Herbert Richards, 370 Riverside Drive, N. Y.

Walter J. Harmer, 100 West 80th Street, N. Y.

Anna E. Lofgren, 575 West 172nd St., N. Y. C.

Mrs. R. A. Wetzel, 218 Tecumseh Avenue, Mt. Vernon, N. Y.

149

Gretchen D. Taylor, 127 Prospect Place, South Orange, N. J.
Mrs. Fitz-Henry Paine, Abington, Conn.

A brief report on the success of the 75th Anniversary Celebra-
tion was given by the chairman of the Celebration Committee, Dr.
Karling, who thanked the members of the institution in the metro-
politan area for their assistance and cooperation in making the
Celebration a success. It was moved by Dr. Rickett that the chair-
man might include this report in a foreword to the issue of TORREYA
covering the Celebration. This was seconded by Dr. Dodge and
passed. 7

With reference to the members of the Club who are now in the
armed forces, Dr. Whaley moved that the Club suspend or impose
a moratorium on their dues so that these members might remain in
good standing for the duration. This was seconded and passed.

The scientific program of the evening then proceeded with re-
ports by several members on their activities during the past sum-
mer.

The meeting was adjourned at 9.40 p.m. The Club then enjoyed
refreshments served by members at the Garden.

Respectfully submitted,

Honor M. HoLtiincHuRST
RECORDING SECRETARY

MINUTES OF THE MEETING OF OcTOBER 21, 1942

The meeting was called to order at 3.30 p.m. by the second
vice-president, Dr. Clyde Chandler, in the Members’ Room
of the Museum Building of the New York Botanical Garden.
Twenty-nine members and friends were present.

The minutes of the preceding meeting were accepted as read.

The following was unanimously elected an associate member:

Rev. James J. Hanlon, 328 West 14th Street, New York City

The scientific program of the afternoon was presented by Dr.
B. O. Dodge who gave an illustrated talk on “Hybrid Vigor or
Heterocaryotic Vigor in the Fungi.” The speaker’s abstract fol-
lows:

Continuation of the work on heterocaryotic vigor has been made possible

by a grant in aid by the American Philosophical Society and by assistance
provided by Dr. W. J. Robbins from private funds advanced for researches

150

on growth substances. It has been previously reported that certain dwarf
races of Neurospora tetrasperma which grow very slowly by themselves
seem to act in a complementary manner to stimulate growth in other rather
slow-growing races, and vice-versa, so that the heterocaryotic mycelia, or
races, grow up to two or three times as rapidly as does either of the individual
components. A rather slow growing race C4, was crossed with a dwarf
race, No. 16, and many ascopores had been isolated at random. Cultures from
these individual ascopores showed that the factors for heterocaryotic vigor
seemed to be heritable. Certain questions arose, however, which indicated
that random selections from dispersed ascospores was not the most desirable
method of procedure. The present work has consisted in the isolation of the
four spores from individual asci, or isolation of the full complement of spores
whenever other than four spores were delimited. All the spores from 131
asci were isolated and grown in culture separately. Of these 118 asci con-
tained four spores, except two or three which contained five spores. Ten
asci contained two normal sized spores and one larger spore. Two asci con-
tained two abnormally large spores and one contained a single giant spore.
In addition, three of the four spores of 39 asci were also isolated and grown
in culture. It was found that 35 of the 118 asci which had four spores
showed that all four spores developed similar cultures which grew vigorously
and all had perithecia. Thirty-five others showed a two and. two pattern in
which two grew vigorously and produced perithecia, while the other two
grew vigorously but very few, if any, perithecia matured. Forty asci showed
a two and two distribution, two cultures growing vigorously, producing an
abundance of ascocarps, while two were dwarfs. These were called double
dwarfs because so far as tested they have shown that two nuclei of both
sexes were present because they fruited with both of the tester strains. The
other asci from which the components were grown showed various sorts of
irregularities which have not as yet been analyzed. In some cases all the spores
were clearly unisexual, as shown by tests.

The advantage in using races of Neurospora tetrasperma for this work over
an obligately heterothallic species such as N. crassa or N. sitophila is that
in the latter forms the nuclei of the opposite sex tend to remain apart even
in mixed cultures so that it is difficult to obtain a heterocaryotic race by
growing two individual unisexual races together in a culture; with JN.
tetrasperma one has no difficulty at all in obtaining heterocaryotic races by
growing two individual races together. In this way it is possible to com-
pare not only the morphological characters exhibited by unisexual or com-
ponent races as compared with a heterocaryotic race composed of the same.
two individual races, but also their comparative growth rates can be ac-
curately measured.

In order to secure fairly accurate growth rates of a large number of in-
dividual races a modification of what we are calling the Beadle and Tatum
tubes are used.

The individual components of 80 bisexual races representing the full in-
heritance of 20 asci have been obtained by plating out conidia, hyphal frag-

Syl

ments, or in the case of the double dwarfs, minute colonies. The growth
rates of a number of bisexual races and of their individual components have
been measured. While this work is only partially completed, there is evi-
dence that the growth rates are probably not determined by single pairs of
factors, although it is clear that such factors exist and that they are in-
herited in a Mendelian fashion. Individual homocaryotic races have been
obtained which show a higher growth rate.

Following the talk, Dr. Mary Schmidt showed some of the ex-
perimental material. The meeting was adjourned at 4:35 p.m. Tea
was then served by friends at the Garden.

Respectfully submitted,

Honor M. HoL_inGHuURST
RECORDING SECRETARY

NEWS NOTES

The Council of the Torrey Botanical Club has decided to pub-
lish the papers presented at the Seventy-fifth Anniversary Celebra-
tion in the 1943 volume of Torreya. This volume will consist
exclusively of these papers and of the Proceedings of the Club.

In furtherance of the effort to conserve quinine and seek for
supplies of cinchona bark from Tropical America, Norman Tay-
lor, the director of Cinchona Products Institute, of New York, is
leaving soon for a survey of plantations and wild sources of bark.
The trip, which includes the region from southern Mexico to
Bolivia, has been authorized by the Board of Commissioners for
the Netherlands East Indies. The chief object is to cooperate’ in
the war effort both with governmental agencies and manufacturers
so that adequate supplies of cinchona bark may be available.

A $1,000.00 fellowship for 1943-1944 is offered by Sigma Delta
Epsilon, the Graduate Women’s Scientific Fraternity. Applica-
tions and reference statements, both in triplicate, should be sub-
mitted before March 1, 1943, to the Fellowship Board.

Women with the equivalent of a Master’s degree, conducting
research in the mathematical, physical or biological sciences, who
need financial assistance to complete their work for the doctorate,

152

and give evidence of high ability and promise are eligible. During
the term of her appointment the appointee must devote the major
part of her time to the approved research project, and not engage
in other work for remuneration (unless such work shall have
received the written approval of the Board before the awarding
of the fellowship or in any later emergency before any new work
shall be undertaken).

Application blanks may be secured from Dr. Eloise Gerry (mark
envelope “Personal” ), care of U. S. Forest Products Laboratory,
Madison, Wisconsin. Announcement of the award will be made
early in April.

THE TORREY BOTANICAL CLUB
OFFICERS FOR 1942

President: C. STUART GAGER
Vice-Presidents: JoHN A. SMALL, F. Treasurer: W. Gorpon WHALEY
CLYDE CHANDLER

‘i Editor: Harotp W. RIcKETT
Recording Secretary: Miss Honor M.

HoLLINGHURST Business Manager: MicHar. LEVINE
Corresponding Secretary: Harotp C. Bibliographer: Mrs. LazELtLa SCHWAR-
Bop TEN

Delegate to the Council, N. Y. Academy of Sciences: W. J. Rossins

Representatives on the Council of the American Association for the
Advancement of Science:

A. E. HitcHcock J. S. Karine
Representative on the Board of Managers of the N. Y. Botanical Garden:
H. A. GLEASoN

Council for 1942

Ex officio members

C. Stuart Gager F. Clyde Chandler Harold W. Rickett
J. S. Karling Harold C. Bold Michael Levine
B. O. Dodge Honor M. Hollinghurst William J. Robbins
John A. Small W. Gordon Whaley
Elected Members

1940-1942 1941-1943 1942-1944
Lela V. Barton Helen M. Trelease Edwin B. Matzke
R. W. Cheney R. C. Benedict Arthur H. Graves
R, A. Harper J. H. Barnhart J. M. Arthur
E. W. Sinnott P. W. Zimmerman W. J. Bonisteel

Committees for 1942
ENDOWMENT COMMITTEE

Clarence Lewis, Chairman J. Ashton Allis
Caroline C. Haynes Helen M. Trelease
Henry de la Montagne

ProcramM CoMMITTEE

eieraidl C. Bold, Chairman (ex officio) Arthur H. Graves
William J. Robbins W. Gordon Whaley
Edwin B. Matzke P. W. Zimmerman

FirLp COMMITTEE
John A. Small, Chairman

Edward J. Alexander H. N. Moldenke Robert Hagelstein
G. G. Nearing Rutherford Platt Michael Levine
Vernon L. Frazee Henry K. Svenson Daniel Smiley, Jr.
Alfred Gunderson Ellys B. Wodehouse Farida A. Wiley
Inez M. Haring Eleanor Friend James Murphy

Loca Frora COMMITTEE
Edwin B. Matzke, Chairman

William J. Bonisteel Harold W. Rickett Dolores Fay

James Edwards Ora B. Smith H. Allan Gleason

John M. Fogg, Jr. Herbert M. Denslow Hester M. Rusk
Cryptogams

Ferns and Fern Allies: R. C. Benedict, W. Herbert Dole, N. E. Pfeiffer
Mosses: E. B. Bartram

Liverworts: A. W. Evans, E. B. Matzke

Freshwater Algae: H.C. Bold

Marine Algae: J. J. Copeland

Fungi: A. H. Graves, J. S. Karling.

Lichens: J. W. Thomson, Jr.

Mysxomycetes: R. Hagelstein

PuBLICATIONS EXCHANGE COMMITTEE
Harold C. Bold, Chairman (ex officio) Amy L. Hepburn Lazella Schwarten

OTHER PUBLICATIONS

OF THE

TORREY BOTANICAL CLUB

(1) BULLETIN

In addition to papers giving the results of research, each issue
contains the INDEx TO AMERICAN BOTANICAL LITERATURE—a very
comprehensive bibliography of current publications in American
botany. Many workers find this an extremely valuable feature of the
BULLETIN.

(2) MEMOIRS

Volume 18, no. 1, 108 pages, 1931, price $2.00. Volume 18, no.
2, 220 pages, 1932, price $4.00. Volume 18 complete, price $5.00.

Volume 19, no. 1, 92 pages, 1937, price $1.50. Volume 19, no.
2, 178 pages, 1938, price $2.00.

(3) INDEX TO AMERICAN BOTANICAL
LITERATURE

Reprinted monthly on cards, and furnished to subscribers at three
cents a card.
Correspondence relating to the above publications should be
addressed to
W. Gorpon WHALEY,
Barnard College,
Columbia University,
New York, N. Y.

TK ol Ge Bes ERE —~ayvr % r
Ran CEN NEW YORI
i

Volume 42 November-December, 1942 Number 620 FAN!

ae O R R FY A GARDEN

A Bi-MontTHLY JouRNAL OF BoTaNnicaAL Notes AND NEws
EDITED FOR
THE TORREY BOTANICAL CLUB
BY

HAROLD A CLUM

John Torrey, 1796-1873

CONTENTS

Some Local Names of Plants—VIII..................00cceees W. L. McATEE 153
Polypetalous Forms of Vaccinium............ W. H. Camp anp C. L. Gitty 168
Carex aestivalis and Carex lurida var. gracilis

on the Glaciated Allegheny Plateau.................. Rosert T. CLAUSEN 174
Papers on the Flora of Alaska—I

The: Genus’ Grcuta yy eo Ns Gases os Reese ete aoe a Fe (oo aahiovon J. P. ANDERSON 176
Some New Forms from the Middle West................ Norman C. Fassett 179
Clarence J. Elting and his Herbarium.................... Homer D. House 181
Book Reviews

Blora of Bukien 205 eee yo ee rene ea rn aN GN R. R. Stewart 190

Itinéraires botaniques dans Vile de Cuba................. Francis E. Liroyp 191

Diary and Travels of the Bartrams...................... Haroitp H. Crum 192

The Carnivorous Plants............... ccc cece e cee ceceees OscaR JANIGER 193
Bields Drips ofthe Club) jp cries oe Be WA Wes RO a ayn et nal da cafes hel teeter nel once oie 196
Proceedings of ‘the @lub! 043i o.b i ig sw SO sers Ca a ee ew LE Gage alt cls eiciere ojstel a4 198
Dates of Publication of TorREYA, VOLUME 42...............2ccceccerecececes 201
| By eK Brel RE a a NL DY AL BIC gO ol Cray Stl iu tera 201
Index to TORREYA, Volume 42...........00 0c cece ec ence cette ects eee eeeees 202

PUBLISHED FOR THE CLUB

By THE FreE Press PRinTING CoMPANY
187 CoLLEecE STREET, BURLINGTON, VERMONT

Entered as second class matter at the post office at Burlington, Vermont,
October 14, 1939, under the Act of March 3, 1879

THE TORREY BOTANICAL CLUB
OFFICERS FOR 1943

President: WILLIAM J. RoBBINS

ist Vice-President: Frep J. SEAVER Recording Secretary: Honor M. Hot-
2nd Vice-President: LELA V. BARTON LINGHURST
Corresponding Secretary: EpwIin B. zreasurer: WW. GoRDON WHALEY
MATZKE Editor: Harotp W. RICKETT
Associate Editors:
Irvine W. BaILey ADRIANCE S. FosTER
Epwarp W. Berry Henry A. GLEASON
STANLEY A. CAIN ARTHUR H. GRAVES
M. A. CHRYSLER JoHN W. SHIVE
Harotp H. Crum R. P. WobdEHOUSE
Business Manager: MicHAEL LEVINE Bibliographer: Mrs. LazELLA SCHWAR-
TEN

Delegate to the Council, N. VY. Academy of Sciences: BERNARD O. DopGE

Representatives on the Council of the American Association for the
Advancement of Science:

JoHn H. BARNHART ALBERT F. BLAKESLEE

Representative on the Board of Managers of the N. Y. Botanical Garden:
Henry A. GLEASON

MEMBERSHIP IN THE TORREY BOTANICAL CLUB

TORREYA

TorREYA was established in 1901 as a bi-monthly publication of the Torrey
Botanical Club for shorter papers and interesting notes on the local flora range of
the Club. It also contains the proceedings of the Club, reports of field trips, and
some book reviews and news notes. The Council of the Torrey Botanical Club has
decided to devote volume 43 of Torreya, 1943, to the publication of the papers pre-
sented in June 1942 at the 75th Anniversary Celebration of the Club, and to the
Proceedings of the Club. This volume will be published in two numbers instead of
the usual six.

ToRREYA is furnished to subscribers in the United States and Canada for one
dollar per year (January-December) : single copies thirty cents. To subscribers
elsewhere, twenty-five cents extra, or the equivalent thereof. Postal or express
money orders, drafts, and personal checks are accepted in payment. Subscriptions
are received only for full volumes.

Of the annual membership dues of the Torrey Botanical Club, $.50 is for a year’s
subscription to TORREYA.

TorREYA is edited for the Torrey Botanical Club by

HAROLD H. CLUM
HunTER Co.Liece, 695 Park AVENUE
New York, N. Y.

TORREYA

VoL. 42 NovEMBER-DECEMBER No. 6

Some Local Names of Plants—VIII *
W. L. McATEE

Correspondents have kindly continued to send local names of
plants, and the writer has been able to glean many in the course of
bibliographic research on birds. Noteworthy accumulations since
the last report are here systematically recorded and alphabetically
indexed. A short list of Literature Cited and suggestions toward
a bibliography of plant vernaculars also are given. The order of
the terms is chiefly that of Heller’s “Catalogue of North American
Plants,” 2nd edit., 1900, and the nomenclature principally that of
Britton and Brown’s “Illustrated Flora,” 2nd edit., 1936.

It may interest readers of Torreya that in 1881, W. R. Gerard,
one of the editors of the Torrey Bulletin, announced an under-
taking to collect and arrange the common names of United States
plants (Amer. Nat. 15:1000).

Literature Cited

Emory, W. H. 1848. Notes of a military reconnoissance from Fort Leaven-
worth, in Missouri to San Diego, in California. 30th Congress, 1st Ses-
sion, Senate Document Ex. 7, 416 pp.

Contains chapters by John Torrey, pp. 135-156, and J. W. Abert, pp.
386-405, the few unusual names in which are here indexed.

Hearne, Samuel. 1911. A journey from Prince of Wales’s Fort in Hudson’s
Bay to the Northern Ocean in the years 1769, 1770, 1771, and 1772
Champlain Society. Toronto, xv+437 pp., illus.

* All of this series have been published in Torreya, No. 1, in Vol. 13:
225-236, 1913; No. 2, 16:235-242, 1916; No. 3, 20:17-27, 1920; No. 4, 26:1-10,
1926; No. 5, 33:81-86, 1933; No. 6, 37:91-103, 1937; and No. 7, 41:43-55,
1941.

TorreEYA for November-December (Vol. 42, 153 to 201) was issued April 24,
1943.

153

154

Relatively few plant names, of which those for four species are cited in
the present glossary.

Lynch, John J. 1942. Louisiana’s state waterfowl refuges. 46 pp. Copies are
on file with the Louisiana Department of Conservation and the U. S.
Fish and Wildlife Service, Washington, D. C.

Contains many local names, both French and English, which for con-
venience are repeated in a 4-page terminal glossary. Terms not pre-
viously recorded in this series nor in the two standard sources men-
tioned are included in the present paper. They are annotated simply as
“Louisiana, Lynch.’ Names not in this compilation, but received in
correspondence from Lynch also are recorded.

Massey, A. B., and R. D. Hatch. 1942. Poisonous plants x x x of Virginia
x x x. Va. Polytechnic Institute, 38 pp. mimeographed.

Richardson, John. 1851. Arctic Searching Expedition: a journal of a boat-
voyage through Ruperts’ Land and the Arctic Sea, etc. 2 vols. Lon-
don.

Contains names of all plants observed and vernacular names for most
of them: Indian, Eskimo, French, and English. Many of the latter are
close to or the same as the modern standard names. Hence only a few
of the most peculiar or interesting terms are cited in the following glos-
sary.

Wied, Maxmilian, Prinz zu. 1839-41. Reise in das innere Nord-America in
den Jahren 1832 bis 1834. Coblenz, 2 vols.

Many German and English and some French and Indian names for
plants. A good proportion are scarcely identifiable. The names here
quoted from this work are noted as, “Weid, Reise, Vol. , p. —.”

As a contribution toward a bibliography of publications dealing
significantly with plant names, the following titles may be cited in
addition to those given in previous installments.

BIBLIOGRAPHY

Ashe, Thomas. 1808. Travels in America, performed in 1806, etc. Contains
lists of medicinal, esculent, ornamental, and useful plants, giving both
Linnean and popular names.

Barton, B.S. 1798. Collections for an essay towards a materia medica of the
United States. Philadelphia.

Bellrose, Frank C. 1941. Duck food plants of the Illinois River Valley.
Ill. Nat. Hist. Survey, Bul. 21(8) Aug.: 235-280, illus. An appendix
(p. 280) lists numerous local names of marsh and aquatic plants.

Boucher, Pierre. 1882. Histoire veritable et naturelle des Moeurs et Pro-
ductions du Pays de la Nouvelle—France. Montreal, iit 164 pp. This
reprint of a work first published in 1663, contains chapters on the woody
plants and on plants cultivated in New France.

Brown, Thomas. 1835. Illustrations of the American Ornithology of Alex-
ander Wilson x x x with x x x representations of the whole sylva of North

155

America. Quarto. London, iii, pp., 124 col. pls. Mtehodical disposition
of the North American sylva, p. iii, has both scientific and vernacular
names, some of the latter unusual.

Carlson, G. G., and V. H. Jones. 1939 (1940). Some notes on the uses of
plants by the Comanche Indians. Papers Mich. Acad. Sci., Arts, and
Letters, 25:517-542. Includes vernacular English and Comanche names.

Carver, J. 1779. Travels through the interior parts of North America in the
years 1766, 1767, and 1768. Chapter 19, pp. 494-526, devoted to trees,
shrubs, roots, herbs, etc., names numerous kinds and recognizably de-
scribes most of them.

Catesby, Mark. 1771. The natural history of Carolina, Florida, and the
Bahama Islands, etc. 2 vols., 220 col. pls. This is a cornerstone of Amer-
ican natural history. It treats plants as numerously as animals. Many of
the vernacular names it employs are still in use and a high proportion of
all are identifiable as the plants are for the most part adequately illus-
trated.

Cooper, J. G. 1859. On the distribution of the forests and trees of North
America, with notes on its physical geography. Ann. Rep. Smithsonian
Inst. for 1858, pp. 246-280. A catalogue in tabular form (pp. 250-266)
includes vernacular names, some of which are not noted in Sudworth’s
“Check List,” 1927.

Coxe, John Redman. 1814. The American dispensatory, etc. 3rd _ edit.
Philadelphia.

Eisenberger, N. F., and G. Lichtensteger. 1750. Piscium, serpentium, insec-
torum x x x guas Marcus Catesby in x x x Carolinae, Floridae x x x
tradidit, etc. 102 pp., 100 col. pls. This volume treats the Catesby material in
the Latin and German languages.

Fernald, M. L. 1910. Notes on the plants of Wineland the Good. Rhodora
12:17-38. Digest of early literature of which numerous titles are cited.
Ganong, W. F. 1910. The identity of the animals and plants mentioned by
the early voyagers to Eastern Canada and Newfoundland. Proc. &
Trans. Roy. Soc. Canada, Ser. 3(3), 1909, Sect. II, pp. 197-242. Assembles

information from about two dozen earlier authors and editors.

Henry, Samuel. 1814. A new and complete American Medical Family
Herbal, etc. New York.

Hitchcock, Edward. 1833. Catalogue of plants growing without cultivation.
Rep. on the Geol.. etc., of Massachusetts, pp. 599-651. Contains many
vernacular names, some exceptional.

Jefferson, Thomas. 1854. Notes on Virginia. The writings of ————,
edited by H. A. Washington. Vol. 8, pp. 281-285. Any of numerous edi-
tions of this work would serve.

Lamb, Wm. H. 1937. Virginia trees. I.—The conifers. Manassas, 112 pp.,

82 figs.

Macoun, John. 1882. Manitoba and the great North-west, etc. Guelph,
xxiit687 pp., illus. Some of the names, particularly of grasses and
sedges, are probably here printed for the first time.

156

McAtee, W. L. 1941. Some local names of Plants—VII. Torreya, 41
(March-April) :43-55. Preceding installment of the present series.

1941. Names of American plants in books on Kalm’s travels.
Torreya, 41 (Sept.-Oct.) :151-160. References to 8 source books, and sys-
tematic list of names in several languages.

Medsger, O. P. 1939. Edible wild plants. xv+323 pp., 19 pls., 80 figs.

Nehrling, Heinrich. 1891. Die nordamerikanische Vogelvelt. xxx+638
pp., 36 col. pls. 10 figs. Contains German vernaculars for numerous
American plants, scientific names for which are given in footnotes.

du Pratz, Le Page. 1758. Histoire de la Louisiane, etc. Paris, 3 vols. The
native plants are chiefly treated in Vol. 2, pp. 1-65 with descriptions and
illustrations sufficient for identification of most of them.

Provancher, L. A. 1862. Flore canadienne; ou description de toutes les
plantes x x x du Canada, etc. Quebec. 2 vols.

Read, Wm. A. 1931. Louisiana-French. Louisiana State University Studies,
No. 5, xxiv+253 pp. Contains numerous French and Indian plant names
and some of other derivations; has a full bibliography and index.

Russell, John L. 1862. [Plants mentioned in Josselyn’s New England’s
Rarities discovered, etc.]. Proc. Essex Inst., 2: 95-115. Identified so far
as practicable.

Saunders, C. F. 1920. Useful wild plants of the United States and Canada.
viiit 275 pp., 69 figs., 20 pls.

Schmitt, Joseph. 1904. Monographie de I’Ile d’Anticosti (Golfe Saint-
Laurent). Botanique, pp. 129-234. Records numerous French names,
some of them provincial.

Seligmann, J. M. and M. Houttuyn. 1772-81. Verzameling van uitlandsche
en zeldzame Vogeln x x x beschreven x x x door G. Edwards en M.
Catesby. Amsterdam, 5 vols. The first volume containing all of the
Catesby material has names of American plants in both “Hoog-” and
“Neder-duitsch.”

Zimmerman, FE. A. W. (Transl. & Editor). 1793. William Bartram’s Reisen
durch Nord- und Sud-Karolina, Georgien, Ost- und West-Florida, etc.
Berlin, xxvit469 pp., 7 pls. The hundreds of plant names in this work
will have to be taken into consideration in any compilation of names of
American plants that aims at completeness.

GLOSSARY

LAMINARIACEAE. 1. Laminaria spp.—Devil’s-apron, C. W. Townsend (Cap-_
tain Cartwright and his Labrador Journal, 1911, p. 257).
SALVINIACEAE. 2. Agolla caroliniana Willdenow.—Water-velvet, Louisiana,

C. Cottam.
EQUISETACEAE. 3. Equisetum hyemale L.—Schachtelhalm (Wied, Reise, 1:
261).

PInacEAE. 4. Pinus banksiana Lambert.—Cyprés of the French voyagers
(Richardson, Arctic Searchings Exp. 2:315, 1851) and of the half-

breeds, Canada (Frank Russell, Explorations in the far North,
1898, p. 103).
5. Pinus virgimana Miller—Yellow pine, John Burroughs (Winter Sun-
shine, 1895 edit., p. 20).
6. Larix laricina Du Roi.—Epinette rouge (French voyagers) ; waggina-
gan (tree that bends; Crees) (Richardson, Arctic Searching Exp.
2:318, 1851) ; “commonly called juniper in Hudson’s Bay.” (Hearne,
Journey, 1911 edit., p. 64).
7. Picea canadensis Miller—Epinette blanche (French voyagers) ; mina-
hik (Crees) (Richardson, Arctic Searching Exp. 2:316, 1851).
8. Tsuga canadensis L.—Canadian fir, Cambria County, Pa., R. M. S.
Jackson (The Mountain, 1860, p. 224).
9. Thuja occidentalis L._—Lebensbaum (Wied, Reise, 2:401).
10. Chamaecyparis thyoides L.—Sweet cedar (Richardson, Arctic Search-
ing Exp. 1:68, 1851).
11. Juniperus horizontalis Moench. (“repens”).—Kriechende Wacholder
(Wied, Reise, 1:389).
12. Juniperus sibirica Burgsdorff—Caw-caw-cue-minick (crowberry)
(Hearne, Journey, 1911 edit., p. 413).
13. Juniperus virginiana L.—Rothe Ceder (Wied, Reise, 1:220).
TypHacEAE. 14. Typha latifolia L.—Cat’s-tail; queue de renard, FEugéne
Bazin (Scenes de la nature dans les Etats-Unis, 1857, 2, p. 144).
15. Typha spp.—Queue de chat, flat rush, jonc plat, jonc matelas, Louisiana,
Lynch.
ZOSTERACEAE. 16. Zostera marina L.—Herbe a languille, Eugéne Bazin
(Scenes de la nature dans les Etats-Unis, 1857, 2, p. 393); herbe
a outarde, Joseph Schmitt (Monographie de 1’Ile d’Anticosti, 1904,
p. 298).
ZANNICHELLIACEAE. 17. Potamogeton foliosus Rafinesque—Gray-duck grass,
herbe canard-gris, Louisiana, Lynch.
18. Potamogeton pectinatus L.—Herbe fine, Louisiana, Lynch.
ALISMACEAE. 19. Sagittaria lancifolia L.—Bull-tongue, langue du _ boeuf,
Louisiana, Lynch.
20. Sagittaria spp—White bull-tongue, Louisiana, C. Cottam.
VALLISNERIACEAE. 21. Vallisneria spiralis L.——Herbe aux canards, Eugéne
Bazin (Scenes de la nature dans les Etats-Unis, 1857, 2, p. 383).
GRAMINEAE. 22. Paspalum distichum L.—Lake grass, Louisiana, Lynch.
23. Panicum dichotomiflorum Michaux.—Sour grass, Allen County, Kan- °
sas, Philip F. Allan.
24. Panicum hemitomon Schultes.—Little cane, canouche, Louisiana, Lynch.
25. Panicum repens L.—Dogtooth grass, dent du chien, Louisiana, Lynch.
26. Panicum virgatum L.—Yellow grass, paille jaune, Louisiana, Lynch.
27. Echinochloa walteri Pursh—Riz de l’ane, riz farouche, riz sauvage,
Louisiana, Lynch.
28. Ziganiopsis miliacea Michaux.—Jonc coupant, Louisiana, Lynch; knife
flag, southeastern Missouri, A. F. Satterthwait (Ecology 2:201,
1921).

158

29. Zizania aquatica L.—Black rice, among a dozen names listed by Chas.
E. Chambliss (Journ. Washington Acad. Sci., 30(5): May 1940)
is additional to those usually recorded; wilde Reiss (Wied,
Reise, 2:83). .

30. Spartina alterniflora Loiseleur-Deslongechamps.—Salt-marsh, coastal
South Carolina, C. Cottam; sea cane, canne du mer, coastal
Louisiana, Lynch.

31. Spartina cynosuroides L.—Cane-marsh, coastal South Carolina, C. Cot-
tam.; hog cane, canne au cochon, quill cane, coastal Louisiana, Lynch.

32. Spartina patens Aiton—Wildcat grass, paille chat-tigre, wire grass,
coastal Louisiana, Lynch.

33. Spartina spartinae Trinius.—Sacahuista, Louisiana, C. Cottam.

34. Bulbilis dactyloides Nuttall—Prairie hay, northern Great Plains, A. A.
Taché (Sketch of n. w. America, 1870, p. 10).

35. Phragmites phragmites L.—Roseau cane, coastal Louisiana, Lynch.

36. Distichlis spicata L.—Paille salé, Louisiana, Lynch.

CYPERACEAE. 37. Scirpus acutus Muhlenberg—Moses weed, New Mexico,
C. Cottam.

38. Scirpus californicus Meyer——Blue grass, Louisiana, C. Cottam; bull-
whip, fouet, jonc rond, round rush, Louisiana, Lynch.

39. Scirpus olneyi A. Gray.—Paille d’oie, jonc au trois quarts, Louisiana,
Lynch. .

40. Scirpus robustus Pursh—Coco, coco grass, Louisiana, C. Cottam,
Lynch; leafy three-square, three-cornered grass, Louisiana; turks-
head, coastal South Carolina, C. Cottam.

41. Cladium jamaicense Crantz.—Redtop, jonc coupant, Louisiana, Lynch.

ARACEAE. 42. Arisaema triphyllum L.—Plant-of-peace, N. N. Puckett (Folk
beliefs of the southern negro, 1926, p. 245).
43. Peltandra glauca Elliott—Cruel man-of-the-woods, N. N. Puckett
(Folk beliefs of the southern negro, 1926, p. 245).
PoNTEDERIACEAE. 44. Pontederia cordata L.—Bull-tongue, langue du boeuf.
Louisiana, Lynch; blue bull-tongue, Louisiana, C. Cottam.
Juncaceaz. 45. Juncus roemerianus Scheele—Jonc negre, jonc piquant,
fouet, whip, Louisiana, Lynch; needle grass, salt rush, coastal
South Carolina, C. Cottam.

LittaceEaAE. 46. Lilium canadense L.—Bitter-root; tra-chin (of the Carrier
Indians), J. K. Lord (The naturalist in x x x British Columbia,
1866, 2, p. 228).

47. Erythronium americanum Ker—Easter lily, Allen County, Kansas,
Philip F. Allan; ‘fawn lily would be better than adder’s tongue.
Still better is the name ‘trout-lily,’ which has recently been pro-
posed,” John Burroughs (Riverby, 1895 edit., p. 25) ; common fawn
lily, Robert B. Troxel (Pennsylvania Game News 13(1) :26, April
1942.

CONVALLARIACEAE. 48. Clintonia borealis Aiton—Bear’s corn, Maine, John
Burroughs (Signs and Seasons, 1895 edit., p. 125) ; Canada may-

159

flower, Richard L. Weaver (New Hampshire Troubador 10(6) :8
Sept. 1940.

TRILLIACEAE. 49, Trillium erectum L.—Red death, P. H. Gosse (Canadian
Naturalist (book), 1840, p. 160).

50. Trillium ovatum Pursh—Herb Paris, Oregon (Richardson, Arctic
Searching Exp. 2:229, 1851.

51. Trillium undulatum Willdenow.—White death, P. H. Gosse (Canadian
Naturalist (book), 1840, p. 160).

JuGLANDACEAE. 52. Juglans nigra L.—Schwarz Wallnussbaum (Wied,

Reise, 1:122).

MyricaceaE. 53. Myrica cerifera L—Wachsbaum (Wied, Reise, 1:171).

SaticaceaE. 54. Salix lucida Muhlenberg —Schmalblattrige Weide (Wied,
Reise, 1:81).

55. Salix purpurea L—Rothe Weide (Wied, Reise, 1:191).

BeETULACEAE. 56. Betula papyrifera Marsh.—Papier-Birke (Wied, Reise,
2:81).

FaGaceakE. 57. Fagus grandifolia Earhart—White, red, mountain, and
water beech, for assumed varieties, Cambria County, Pa. R. M. S.
Jackson (The Mountain, 1860, p. 221).

58. Quercus phellos L.—Weideneiche, weiden-blatterige- Eiche (Wied,
Reise, 2:377, 1:145).

59. Quercus prinus L.—Kastanien-Eiche (Wied, Reise, 1:56).

60. Quercus stellata WWangenheim.—Spalt oak, Geo. H. Cook (Geology of
Cape May, N. J., 1867, p. 75).

UtmaceaE. 61. Ulmus thomasi Sargent (“suberosa”).—Wahu-Ulme (Wied,
Reise, 1:309).

MoraceEak. 62. Papyrius papyrifera L.—Papier-Maulbeerbaum (Wied, Reise,
1:40).

ARISTOLOCHIACEAE. 63. Aristolochia serpentaria L.—Schlangenwurzel (Wied,
Reise, 1:75).

PoLYGONACEAE. 64. Polygonum spp.—Curage, Louisiana, Lynch.

CHENOPODIACEAE. 65. Salicornia spp.—Baloney-grass, salt-grass, San Luis
Obispo, California, C. Cottam; sea-fennel, Thos. F. De Voe (The
Market Assistant, 1867, p. 365). ;

66. Salsola kali L.—Tumbleweed, Texas, V. W. Lehmann (Wildlife Re-
view 22:41, 1939).

AMARANTHACEAE. 67. Acnida alabamensis Standley—Chou gras, Louisiana,
Lynch.

PHYTOLACCACEAE. 68. Phytolacca decandra L.—Crow-berry, chou gras, red-
ink berry, Doris M. Cochran (Nature Mag. 35(2) :74, Feb. 1942;
Kermesbeere (Wied, Reise, 1:33).

NyctTaGINAceaE. 69. Allionia nyctaginea Michaux.—Snotweed, Allen
County, Kansas, Philip F. Allan.

NeELUMBONACEAE, 70. Nelumbo lutea Willdenow.—Big bonnet, Mississippi,
C. Cottam; lily-nut, Maurice Thompson (Byways and Bird Notes,
1885, p. 103).

160

CapoMBACEAE. 71. Brasenia schreberi Gmelin—Small bonnet, Mississippi
C. Cottam.
NyMPHAEACEAE. 72. Castalia elegans Hooker.—Pagayeur, Louisiana, Lynch.
73. Castalia flava Leitner —Herbe au coeur, Louisiana, Lynch.
74. Castalia odorata Dryander—Beaver root, J. G. Millais (Newfound-
land and its untrodden ways, 1907, p. 236); pagayeur, Louisiana,
Lynch.
75. Nymphaea advena Solander—Can-dock, splatter-dock, Philadelphia,
Pa., B. S. Barton (A discourse on x x x Nat. Hist., 1807, p. 48).
MacGnoiiaceak. 76. Magnolia virginiana L—Spoonwood, Geo. H. Cook
(Geology of Cape May, N. J., 1867, p. 76).
77. Liriodendron tulipifera L—Tulpenbaum (Wied, Reise, 1:48; wild
poplar, Cambria County, Pa., R. M. S. Jackson (The Mountain,
1860, p. 227).
78. Illictum floridanum Ellis —Aniseed tree, H. B. Croom (Amer. Journ.
Sci. & Arts 26:319, 1834).
ANNONACEAE. 79. Asimina triloba L.—Hoosier banana, Indiana.
RANUNCULACEAE. 80. Actaea alba L.—Racine d’ours (French Canadians) ;
musqua-mitsu-in (bear’s food, Crees) (Richardson, Arctic Searching
Exp. 1:82, 1851).
81. Aconitum napellus L.—Queen’s fettle, Great St. Lawrence, Nfd., J. G.
Millais (Newfoundland and its untrodden ways, 1907, p. 144).
82. Hepatica hepatica 1.—Red coon-root, N. N. Puckett (Folk beliefs of
the southern negro, 1926, p. 375).
83. Pulsatilla patens L.—Rothe Kalbsblume (translation of an Indian
name meaning red calf-flower) (Wied, Reise, 2:314).
LauRACEAE. 84. Benzgoin aestivale L—Gewurzholz (Wied, Reise, 1:223).
FuMARIACEAE. 85. Bicuculla cucullaria L.—Staggerweed, Virginia (Massey
and Hatch, 1942, p. 6). The authors record this name as being ap-
plied also to Bicuculla canadensis Goldie, Capnoides flavulum
Rafinesque, and Delphiniwm tricorne Michaux.
CrucIFERAE. 86. Draba caroliniana Walter—Shad-blossom, Philadelphia,
Pa., B. S. Barton (A discourse on x x x Nat. Hist., 1807, p. 28).
HypraNGEACEAE. 87. Hydrangea quercifolia Bartram.—Swamp snow-ball,
B. L. C. Wailes (Rep. Agr. Geol. Miss., 1854, p. 344).
ALTINGIACEAE. 88. Liquidambar styraciflua L—Storaxbaum (Wied, Reise,
Legit).
HAMAMELIDACEAE. 89. Hamamelis virginiana L.—Zauberhaselnuss (Wied,
Reise, 2:343).
PLATANACEAE. 90. Platanus occidentalis L.—Wasser-Ahorn, Wasser-Buche,
Germans in Pennsylvania (Wied, Reise, 1:72).
RosacgEaE. 91. Spiraea latifolia Aiton—Wiedenblatterige Spierstande (Wied,
Reise, 1:81).
92. Rubus chamaemorus L.—Bethago-tominick (Crees); dewater berry
(Hearne, Journey, 1911 edit., p. 411).
93. Rubus deliciosus James.—False raspberry (Colorado Agr. Exp. Sta.
Bul. 445:35, 1938). Commenting on the book name Boulder rasp-

161

berry, the late Francis Ramaley wrote me (Feb. 21, 1941), “I have
lived in Boulder for forty years and never heard this bush called
anything but ‘Thimbleberry’—never the word ‘Boulder’ attached to
it.”

94. Rubus lasiococcus Gray.—Fuzzy mountain-dewberry, Oregon, Helen
M. Gilkey.

95. Rubus nivalis Douglas —Small mountain-blackberry, Oregon, Helen M.
Gilkey.

96. Rubus pedatus Smith—Red mountain-dewberry, Oregon, Helen M.
Gilkey.

MatacesE. 97. Malus fusca Rafinesque (“rivularis”).—Powitch tree (Rich-
ardson, Arctic Searching Exp. 2:294, 1851). This is from the
Chinook, pauitsh.

-98. Amelanchier canadensis L.—Bois de fléche, French voyagers (Richard-
son, Arctic Searching Exp. 2:294, 1851).

AMYGDALACEAE. 99. Laurocerasus caroliniana Miller—Lauria mundi,
B. L. C. Wailes (Rep. Agr. Geol. Miss., 1854, p. 342).

100. Padus virginiana L.—Traubenkirsch (Wied, Reise, 1:291).

Mimosaceae. 101. Morongia uncinata Willdenow.—Saw-brier, Thomas Nut-
tall (Travels into the Arkansa Territory, 1821, p. 180).

CAESALPINACEAE. 102. Cercis canadensis L.—Shad-Blossom, Philadelphia,
Pa., B. S. Barton (A discourse on x x x Nat. Hist., 1807, p. 28).

103. Gymnocladus dioica L.—Bonduc, Edwin James (Long’s Exp. Rocky
Mts., Thwaites edit., 1905, Pt. 1, p. 213).

104. Hoffmannseggia sp—Chufa (from the root nodules), mesquite weed,
Hansford County, Texas, Philip F. Allan.

Fapaceakz. 105. Baptisia leucantha Torrey and Gray.—Prairie indigo, J. W.
Abert (in Emory, W. H., Military Reconnaissance, 1848, p. 399).

106. Psorolea esculenta Pursh.—Wild turnip (Wied, Reise, 1:321).

107. Amorpha fruticosa L.—Pride-of-Barbadoes, B. L. C. Wailes (Rep. Agr.
Geol. Miss., 1854, p. 343).

108. Parosela dalea L—Woods clover, Allen County, Kansas, Philip F.
Allan.

109. Sesban macrocarpa Muhlenberg.—Indigo, acacie, Louisiana, Lynch.

110. Daubentonia drummondiu Rydberg —Coffee bean, Louisiana, Lynch.
This term and coffee weed are applied in various parts of the South
to almost any conspicuous wild legume.

111. Astragalus emoryanus Rydberg.—Red-stemmed peavine, Texas, Frank
P. Matthews (Journ. Amer. Veterinary Med. Assoc. 97:125, 1940).

112. Meibomia sp—Wood sage, Allen County, Kansas, Philip F. Allan.

113. Alysicarpus vaginalis L.—Alice clover, Herbert L. Stoddard (6th Ann.
Rep. Cooperative Quail Study Assoc., 1938, p. 10).

114. Lespedeza striata Thunberg.—Buffalo, Carolina, China, Georgia, and
oldfield, clover, southeastern States, J. W. Kistler (N. C. Wildlife
Conservation 4(12):5, Dec. 1940).

115. Vicia angustifolia L—Augusta vetch, Herbert L. Stoddard (7th Ann.
Rep. Cooperative Quail Study Assoc., 1939, p. 16).

162

116. Lathyrus hirsutus L—Wild winter-pea, Herbert L. Stoddard (7th Ann.
Rep. Cooperative Quail Study Assoc., 1939, p. 18).

ZYGOPHYLLACEAE. 117. Covillea tridentata De Candolle—lIodeodondo of the
Mexicans, John Torrey (in Emory W. H., Military Reconnoissance,
1848, p. 138).

Me iacesE. 118. Melia azedarach L.—Bead tree, John Latham (Gen. Hist.
Birds, 5, 1822, p. 145).

EUPHORBIACEAE. 119. Croton capitatus Michaux—Bighead doveweed, Okla-
homa, Verne Davison (Wildlife Review 22:38, 1939) ; billy-goat
weed, hogwort, Herbert L. Stoddard (3rd Ann. Rep. Cooperative
Quail Study Assoc., 1936, p. 14).

120. Croton texensis Klotzsch—Texas doveweed, Verne Davison (Wild-
life Review 22:38, 1939).

121. Euphorbia lathyrus L—Sassy Jack, mountains of Virginia (Massey
and Hatch, 1942, p. 5). ;

EMPETRACEAE. 122. Empetrum nigrum .—Black-berried heath, John
Latham (Gen. Hist. Birds, 10, 1824, p. 261); black crake-berry,
G. G. Macdougall, Transl. (Graah, W. A., Narrative of an Expe-
dition to the east coast of Greenland, 1837, p. 135) ; nischa-minnick
(Gray-goose berry, Crees), Hearne (Journey, 1911, p. 411).

123. Ceratiola ericoides Michaux.—Sand hill rosemary, H. B. Croom (Amer.
Journ. Sci. & Arts 26:315, 1834).

ANACARDIACEAE. 124. Schinus terebinthifolius Raddi—Brazilian or Mexican

pepper-tree, Florida holly, Vero Beach, Florida.
125. Rhus hirta L.—Hirschkolbenbaum (Wied, Reise, 2:18).
126. Rhus trilobata Nuttall—Squaw berry, southern Utah, C. Cottam.

CyRILLACEAE. 127. Cliftonia monophylla Lamarck—Buck-wheat tree, H. B.
Croom (Amer. Journ. Sci. & Arts 26:319, 1834).

ILacacEAE. 128. Ilex cassine L—Yapa shrub, John Latham (Gen. Hist.
Birds, 5, 1822, p. 142).

129. Ilex decidua Walter—Swamp spice, B. L. C. Wailes (Rep. Agr. Geol.
Miss., 1854, p. 344).

STAPHYLEACEAE. 130. Staphylea trifolia L.—Dreiblatterige Pimpernuss
(Wied, Reise, 1:294).

ACERACEAE. 131. Acer negundo L.—Manitoba maple, Ottawa, Canada.

Viraceaz. 132. Vitis cordifolia Michaux.—Choke grape, B. L. C. Wailes
(Rep. Agr. Geol. Miss., 1854, p. 346).

TrtacEaAE. 133. Tilia heterophylla Ventenat (“grandifolia’”).—Gross-
blatterige Linde (Wied, Reise, 1:145).

Matvaceae. 134. Hibiscus grandiflorus Michaux.—Cotton rose, B. L. C. -
Wailes (Rep. Agr. Geol. Miss., 1854, p. 346).

TAMARICACEAE. 135. Tamarix gallica L.—Salt cedar, coastal Louisiana,
Lynch.

136. Fouquiera splendens Engelmann.—Boojum tree, Superior, Arizona,
H. K. Gloyd (Chicago Naturalist 3(3) :73, Oct., 1940).
PASSIFLORACEAE. 137. Passiflora incarnata L.—Apricot, the fruit or maypop,

Margaret W. Morley (The Carolina Mountains, 1913, p. 68).

163

CactacesE. 138. Opuntia bigelovii Engelmann.—Teddy-bear cholla, Su-
perior, Arizona, H. K. Gloyd (Chicago Naturalist 3(3): 71, Oct.,
1940).

139. Opuntia fulgida Engelmann.—Jumping cholla, Superior, Arizona, H. K.
Gloyd (Chicago Naturalist 3(3) :71, Oct., 1940).

140. Opuntia polyacantha Haworth (“glomerata”)—Crapaud vert, French
voyagers (Richardson, Arctic Searching Exp. 2:279, 1851).
ELAEAGNACEAE. 141. Elaeagnus argenta Pursh.—Stinking willow (fur trad-
ers); Tap-pah (gray berry, Chepewyans) (Richardson, Arctic
Searching Exp. 1:145, 1851) ; silvery oleaster (ibid. p. 199) ; napow-

muskwaniman (white bear-berry, Crees) (Op. cit. 2:307, 1851).

142. Lepargyraea argentea Nuttall—Wied (Reise, 2:80) wrote “Graines
de boeuf,” considering “graisse de boeuf” an error; he was mistaken,
however, as the name “beef-suet tree’ indicates.

OnaGRACEAE. 143. Chamaenerion latifolium L.—Indian wickup, P. H. Gosse
(Canadian Naturalist (book), 1840, p. 298).

ARALIACEAE. 144. Aralia racemosa L.—King-of-the-woods, N. N. Puckett
(Folk beliefs of the southern negro, 1926, p. 246).

145. Aralia spinosa L.—Devil’s-club, Cambria County, Pa., R. M. S. Jack-
son (The Mountain, 1860, p. 237).

AmmtacEAk. 146. Hydrocotyle ranunculoides L. £—Water parsley, parasol,
Louisiana, Lynch.

147. Cicuta maculata L.—California fern, Virginia (Massey and Hatch,
1942, p. 18) ; carrotte de Moreau (after a man who died from eat-
ing the root), manito-skatask, Crees (Richardson, Arctic Searching
Exp. 1:95, 1851).

148. Sium cicutaefolium Schrank.—Queue de rat, French Canadians;
uskotask, Crees (Richardson, Arctic Searching Exp. 1:95, 1851).

149. Heracleum lanatum Michaux.—Alexander, C. W. Townsend (Captain
Cartwright and his Laborador Journal, 1911, p. 82).

CorNACEAE. 150. Cornus amomum Miller —Hartriegel (Wied, Reise, 1:326).

151. Cornus stolonifera Michaux.—Osier rouge (Richardson, Arctic Search-
ing Exp. 2:273, 1851).

EricacEaE. 152. Dendrium buxifolium Berg.—Heather, Margaret W. Mor-
ley (The Carolina Mountains, 1913, p. 253).

153. Ledum groenlandicum Oeder.—Indian tea, P. H. Gosse (Canadian
Naturalist (book), 1840, p. 300).

154. Kalmia latifolia L—Ivy, Margaret W. Morley (The Carolina Moun-
tains, 1913, p. 56).

155. Xolisma ligustrina L.—Male berry, staggerbush, Virginia (Massey and
Hatch, 1942, p. 15). The authors note that the second name is used
also for Leucothoé catesbaei Walter.

156. Gaultheria humifusa Graham.—Mountain wintergreen, Oregon, Helen
M. Gilkey. :

157. Gaultheria procumbens L.—Pine ivy, Thos. F. De Voe (The Market
Assistant, 1867, p. 394).

164

158. Uva-urst uva-urst L—Graine d’ours (bear-berry), sac a commis, Bear
Lake, Canada, George Keith (in Masson, L. R., Les Bourgeois de la
Compagnie du Nord-QOuest, 2, 1890, p. 102) ; sakakomi, Sakkakomi-
Pflanze (Wied, Reise, 2:81, and 1:445).

159. Vaccinium ovalifolium Smith—Tall blue huckleberry, Oregon, Helen
M. Gilkey.

160. Vaccinium ovatum Pursh—Evergreen huckle-berry, Oregon, Helen M.
Gilkey.

161. Vaccinium parvifolium Smith—Peacock berry. W. L. Dawson (Birds
ot Washington, 2, 1909, p. 577) ; red huckleberry, Oregon, Helen
M. Gilkey.

162. Vaccinium uliginosum L—Ground whortle (old name), whorts (new
name), C. W. Townsend (Captain Cartwright and his Labrador
Journal, 1911, p. 34).

SAPOTACEAE. 163. Bumelia tenax L—Sloe berry, coastal Georgia, C. Cot-
tam.

EBENACEAE. 164. Diospyros virginiana L.—American medlar, Thos. F. De
Voe (The Market Assistant, 1867, p. 386).

OLEACEAE. 165. Forsythia sp—*‘Sunshine bush, it deserves to be called,”
Bradford Torrey (Clerk of the Woods, 1903, p. 2).

166. Forestiera neo-mexicana A. Gray—Wild olive, New Mexico, A. E.
Borell.

GENTIANACEAE. 167. Frasera carolinensis Walter (“waltheri’)—Falsche
Colombo-wurzel (Wied, Reise, 1:170).

APOCYNACEAE. 168. Apocynum androsaenifolium L—Angel’s turnip, N. N.
Puckett (Folk beliefs of the southern negro,* 1926, p. 245) ; herb
a la puce (from its irritating effects) ; this name applied also to
A. sibiricum Jacquin (hypericifolium Aiton) (Richardson, Arctic
Searching Exp. 1:121, 1851).

CONVOLVULACEAE. 169. Convolvulus arvensis L.—Possession vine, Texas Pan-
handle, Philip F. Allan; tie vine, B. L. C. Wailes (Rep. Agr. Geol.
Miss., 1854, p. 344).

LapraTae. 170. Mentha spicata L—Green mint, Thos. F. De Voe (The Mar-
ket Assistant, 1867, p. 364).

SoLANCEAE. 171. Solanum carolinense L.—Tread saft, N. N. Puckett (Folk
beliefs of the southern negro, 1926, p. 246).

172. Lycopersicon lycopersicon L.—Liebesapiel (Wied, Reise, 1:191).

173. Datura stramonium L—Stechapfel (Wied, Reise, 1:33).

174. Nicotiana quadrivalvis Pursh—Manascha (Mandan Indians), Tabacks-
pflanze (Wied, Reise, 2:90 and 122).

BIGNONIACEAE. 175. Catalpa catalpa .—Petalira (“which as well as catalpa,
the received appellation, may be a corruption from Catawba, the
name of the tribe by whom x x x the tree may have been intro-

* Some obvious misidentifications in this book have been excluded, and pos-
sibly the records under Nos. 43 and 168 also should have been rejected.

165

duced’), Smithland, Kentucky. Edwin James (in Long’s Exp. to
Rocky Mts., Thwaits edit., 1905, Part I, p. 84).

MartyNiacEAE. 176. Martynia louisiana Miller—Cuckold’s horns, Edwin
James (in Long’s Exp. to the Rocky Mts., Thwaites edit., 1905, Part
2, p. 44).

CAPRIFOLIACEAE. 177. Viburnum lentago L—Partridge berry, Thos. F. De
Voe (The Market Assistant, 1867, p. 384).

178. Viburnum opulus L.—Mongs6-a mina (moose-berry, Crees) ; dunne-
ki-e (Indian berry, Dog-rib and Hare Indians) (Richardson, Arctic
Searching Exp. 1:120 and 2:298, 1851).

179. Viburnum pauciflorum Pylaie—Pembina, French voyagers; nipi-minan
(water-berry, Crees) (Richardson, Arctic Searching Exp. 1:120,
1851, and 2:298). This allocation of the term pembina, is more
probably correct than that quoted from Clapin under No. 157 in the
preceding installment.

CucurBITACEAE. 180. Pepo foetidissima Humbolt, Bonpland, and Kunth.—
Prairie gourd, J. W. Abert (in Emory, W. H., Military Recon-
noissance, 1848, p. 398).

CicuoriaceEazE. 181. Nabalus albus L. (“rubicunda”)—Lowenherz (Wied,
Reise, 1:75).

182. Nabalus serpentarius Pursh.—Gall-of-the-earth, B. L. C. Wailes (Rep.
Agr. Geol. Miss., p. 346).

AMBROSIACEAE. 183. Ambrosia elatior L.—Short ragweed, Oklahoma, Verne
Davison (Wildlife Review 22:38, 1939); Georgia, Herbert L.
Stoddard (op. cit. p. 43).

184. Xanthium sp—Cuckold bur, Thomas Nuttall (Travels into the
Arkansa Territory, 1821, p. 58).

ComposiTaE, 185. Laciniaria pycnostachya Michaux.—Kansas gayfeather,
Allen County, Kansas, Philip F. Allan; pinette de prairie, J. W.
Albert (in Emory, W. H., Military Reconnoissance, 1848, p. 398).
Should be epinette.

186. Gutierrezia spp—Fireweed, lightning-brush, Utah, C. Cottam.

187. Grindelia squamosa Pursh—Epinette de prairie (Wied, Reise, p. 517).
See 185; while these authorities differ, both may have correctly
recorded usage.

188. Heterotheca grandiflora Nuttall—Telegraph weed, Santa Cruz County,
Calif., A. C. Hawbecker (Journ. Mammalogy 21(4) :389, 1940).

189. Heterotheca subaxillaris Lamarck—Camphor-weed, Texas, V. W.
Lehmann (Wildlife Review 22:41, 1939).

190. Baccharis halimifolia L—Manglier; mung, Louisiana, Lynch. These
terms are reminiscent of names applied to Iva. (See this series, 1,
1913, No. 102.) Evidently there is popular confusion of the two
genera. Sand myrtle, coastal South Carolina, C. Cottam.

191. Acanthospermum hispidum De Candolle——Texas spur, star bur (News
Letter, Cooperative Quail Study Assoc., Thomasville, Ga., 3, Dec.,
1942, p. 6).

192.

166

sas, Philip F. Allan.

193.

Silphium perfoliatum De Candolle—Pitcher-plant, Allen County, Kan-

Echinacea purpurea L.—Rattlesnake weed, J. W. Abert (in Emory,

W. H., Military Reconnoissance, 1848, p. 387).

Texas Panhandle,

Philip F.

a corruption of vanilla),
1854, p. 342).

Clover, Buffalo ....... 114
Clover, Carolina ...... 114
Clover Chinaleeeeeee 114
Clover, Georgia ...... 114
Clover, Oldfield ...... 114
Clover, Woods ...... 108
Coco 2G. skelnectrs.csccis 40
Coco) grass) ss aoeeeee ee 40
Coffeewhean¥enaene eee 110
Coffee weed ......... 110
Colombo wurzel, Falsche 167
Common fawn lily .... 47
Coon-root, Red ...... 82
(CROMANSAA? soocooccoe 68
Corn} Beancsireee eee 48
Cottontxose) see eee 133
Crakeberry, Black .... 122
Grapaudiverte see 140
Crenate milkweed 197
Crowberryeeeeo eee 12
Cruel man-of-the-woods 43
Guckoldibursseseee eee 184
Cuckold’s horns ...... 176
C@urage! is. cease lees 64
G@yipres! sites cecoose 4
Dent du chien ....... 25
Devil’s-apron .........- 1
Devil7s-clubseeeeeeeee = 145
Dewater berry ....... 92
Dock i€ant ash oaueeces 75
Dock Splatter sao 75
Dogtooth grass ....... 25
Doveweed, Bighead.... 119
Doveweed, Texas 120
Dreiblatterige Pimper-

TUS racustevenodsnsyenciohelate 130
Dunneé-ki-e .........- 178
Hastermlily meets 47
Eiche,: Kastanien ..... 59
Eiche, Weiden ....... 59

Eiche, Weiden-blat-
CERISE” Was etccte ers 58
Epinette blanche ..... a

Epinette de prairie 185, 187
6

194. Borrichia frutescens L.—Button weed, coastal South Carolina, C.
Cottam.
195. Gaillardia sp—Indian-blanket flower,
Allan.
196. Artemisia gnaphalodes Nuttall—Wermuth (Wied, Reise, 1:556).
197. Erechtites hweracifolia L—Crenate milkweed, P. H. Gosse (Canadian
Naturalist (book), 1840, p. 288).
198. Synosma suaveolens L.—Vinella (doubtless
B. L. C. Wailes (Rep. Agr. Geol. Miss.,
INDEX
INCACI OW ria Py syetecenenereuene 109) Blueverassy cee 38
Ahorn, Wasser ....... 90 Blume, Rothe Kalbs .. 83
Alexander etisrs aiysis ors 149 Bois de fléche ....... 98
Alice clover .......... 1G OAC soccoosessece 103
American medlar ..... 164 Bonnet, Big ........- 70
Angel’s turnip ........ 168 Bonnet, Small ....... 71
Aniseed tree ......... 7o) JBoojumetreenny ysis 137
Apfel, Liebes ......... 172 Boulder raspberry .... 93
Nal, SEIN Scocq80cn 173 Brazilian pepper-tree .. 124
IA TICODe em 137. Brier, Saw .......... 101
Augusta vetch ....... 115 Brush, Lightning 186
Baloney-grass ........ 65 Buche, Wasser ...... 90
Banana, Hoosier ..... 79 Buck-wheat tree ...... 127
Baum, Storax ........ gg Buffalo clover ........ 114
Baum, Tulpen ........ 77 Bull-tongue ........ 19) 44
Beaditrec eee eerie 11g Bull-tongue, Blue 44
Tea, COIS sooscnse- 119 Bull-tongue, White 20
Bear-berry, White .... 141 Bull-whip ............ 38
Bee's. GORD oo 50005000 4g Bur, Cuckold ........ 184
IRGEREIP TOG Soocccoodc 74 Bur, Star ...-..--.--- 191
Beech, Mountain ..... 57 Bush, Sunshine ...... 165
Beech Red eee 57 Button weed ......... 194
Beech, Water ........ 57 California fern ....... 147
Beech, White ........ 57 Camphor weed ....... 189
Beef-suet tree .....-. 142 Canada Mayflower .... 48
IeSie, INSHMES sosccas 6g Canadian fir ......... 8
Berry, Crow ...... 12, 68 Can-dock ............ 75
Berry, Dewater ...... 92 Cane, Hog eee eet ewes 31
Berry, Gray We A ea oe 141 Cane, Little eee ee eee 24
Berry, Grey-goose .... 122 Cane, Quill .......... 31
Bernyneindianweee eet 17g Cane, Roseau ......-. 35
lean WENS sosonccace 155 Gane; (Sea Sa.coe teers 30
Berry, Moose ........ 17 CamMesnersn osooaossoc 31
Berry, Partridge ..... 177. Canne au cochon ..... 31
Berry, Peacock ...... 161 Canne du mer ....... 30
Berry, Red-ink ....... 3 CEMORON ssscoosuo0ccs 24
1ESaAry SID Sonooeabec 163u).Garolinaycloversuse see 114
Inca, SOE? Goccocec 126 Carrotte de Moreau 147
Berry, Thimble. . OR Cael cavgsscessesc 175
IBEGRYARWateGEeeeeeeee 170), (Cates:taill | Sterne cr. 14
Bethago-tominick ..... 92 Caw-caw-cue-minick ... 12
lites [SOaEE Goacacc-so CK) Crake, Selle soéocoosoe 135
Bighead doveweed .... 119 Cedar, Sweet ........ 10
Billy-goat weed ...... ie) (Geceie, INGHNS s5655000c 13
Birke, Papier ........ S56) Chinaiclovemuacreres ce 114
Bitterroot saeco. 460) Chokeverapeseeeeeeeee 132
Black-berried heath 122 Cholla, Jumping ...... 139
Black crakeberry ..... 122 Cholla, Teddy-bear .... 138
IGE THES acocosanobas- 20 ee Chou eras) eae ce , 68
Blossom, Shad CG, 02 Chitnky egeaoacsocdsocs 104
Blue bull-tongue ...... 44 Clover, Alice ........ 113

Epinette rouge .......

Evergreen huckleberry. 160
Falsche Colombo-wurzel 167
False raspberry ...... 93
IDeiyan WIRG coco: caccce 47
Fawn lily, Common 47
Ikan SEY Goooocasc 65

Fern, California ...... 147
laiie, (Carncberm soocooc 8
Gem weed erento 186
Flag, Knife ......... 28
IMENE TAIN soococccco00 15
Florida holly ......... 124
Flower, Indian-blanket. 195
INOTe tin ieeeeerererlercr sie 38, 45
Fuzzy mountain-dew-

Derry eine eae S elelelevereiels 94
Gall-of-the-earth ..... 182
Gayfeather, Kansas.... 185
Georgia clover ....... 114
Gewurzholz .......... 84
Gourd, Prairie ....... 180
Graine d’ours ........ 158
Graines de boeuf 142
Graisse de boeuf ..... 142
Grape, Choke ........ 132
Grass, Baloney ...... 65
Gees, Bile sooc000006 38
Grass, (Cae scoocadcoc 40
Grass, Dogtooth ...... 25
Grass, Gray-duck 17
(Grassheakeererneriarr 22
Grass, Needle ........ 45
Grass, Selle soocoocsoc 65
(Grass, Some sodcccooce 23
Grass, Three-cornered . 40
Grass, Wildcat ...... 32
Grass; Wire ......... 32
Grass, Yellow ........ 26
Greeny WAY oobdooc006 141
Gray-duck grass ...... 17
Gray-goose berry ...... 122
Green mint .......... 170
Grossblatterige Linde.. 133
Ground whortle ...... 162
Hartriegel ........... 150

Haselnuss, Zauber .... 89

Heath, Black-berried .. 122
ISIGAES? 5000000000000 152
Herb a la puce ...... 168
Herb Paris .......... 50
Herbe a l’anguille .... 16
Herbe a outarde ...... 16
Herbe au coeur ...... 73
Herbe aux canards ... 21
Herbe canard-gris .... 17
Herbe fine ..........- 18
Hirschkolbenbaum 125
laloye GINS Soagaceoouood 31
los wonthieeceirocier 119
Holly, Florida ....... 124
Hoosier banana ...... 79
Huckleberry, Evergreen 160
Huckleberry, Red .. 161
Huckleberry, Tall bikie. 159
Indian, berry ........ 178
Indian tea ........... 153
Indian wickup ....... 143
Indian-blanket flower .. 195
ari SON ace cosbeteneiaiees love 109
Indigo, Prairie ...... 105
Iodeodondo .......... 117
lai fec Geen Clos yoo ce 154
IBAR, IBM “Gooosdooace 157
Jone au trois quarts .. 39
Jone coupant ...... 28, 41
Jone matelas ......... 15
Jonecsnegre esas ee 45
Jonchpiquants eee 45
Jonchplatena-ooence 15
Ore Rol ooocsocoacs 38
Jumping cholla ....... 139
Jainiperd Sicsic-eencearnes cue 6
Kalbsblume, Rothe .... 83

167

Kansas gayfeather .... 185
Kastanien-Eiche ...... 59
Kermesbeere ......... 68
King-of-the-woods 144
Kirsch, Trauben ...... 100
Kaine WE? so 5500008 28
Kriechende Wacholder 11
ILAIKS FRFASS sudaooacus 22

Langue du boeuf ....19, 44
Lauria mundi 99
Leafy three-square .... 40

Bebensbaumi eae 9
Liebesapfel .......... 172
Lightning brush ...... 186
Itsy, WARE Gooooebos 47
ilyaHaw nee een 47
ILM IDROGKE “Go oodocuss 47
Ibithy, “ARROBIE oo ndoeoo000 47
Linde, Grossblatterige.. 133
ittlencanemmanmcee ccs 24
Lowenherz ........... 181
Male berry .......... 155
Manaschalen anne: 174
Mewes oogoccoscu0s 190
Manito-skatask ....... 147
Manitoba maple ...... 131
Man-of-the-woods, Cruel 43
Maple, Manitoba 131

Maulbeerbaum, Papier. 62

Mayflower, Canada ... 48
MERRION) SooodcadeouGs 137
Medlar, American .... 164
Mesquite weed ....... 104
Mexican pepper-tree 124
Milkweed, Crenate .... 197
MbbagA I Go dc00000000 7
Mint, Green ......... 170
Mongs6-a mina ...... 178
Moose berry ......... 178
Moses weed .......... 37
Mountain beech ...... 57
Mountain-blackberry,

Small vaactseesmeeeiare 95
Mountain-dewberry,

IDSEZAZ Golceoco Danas 94

Mountain-dewberry, Red 96

Mountain-wintergreen . 156
Webs emes oreo dose bom mae 190
Musqua-mitsu-in ..... 80
Myrtle, Sand ........ 190
Napow-muskwa-minan ..141
Needle grass ......... 45
Nipi-minan .......... 179
Nishca-minnick ....... 122
INE, ILA? Gooocoo0a0s 70
Oak Spalt eerie eie. 60
Oldfield clover ....... 114
Oleaster, Silvery ..... 141
Olive, Wild ......... 166
Osier rouge ......... 151
Ragayeunmenrreccn cir 72, 74
Paille chat-tigre ...... 32
Paille d’oie .......... 39
Paille jaune ......... 26
Raillemsalemercemere cee 36
Rawitshaneeeeceeccoer 97
Papier-Birke ......... 56
Papier-maulbeerbaum . 62
Parasol Wyse sts 146
Parsley, Water ...... 146
Partridge berry ...... 177
Peacock berry ........ 161
Peavine, Red-stemmed . 111
Rembinawaneeenecert 179
Pepper-tree, Brazilian . 124
Pepper-tree, Mexican 124
Petalinamen coerce 175

Pimpernuss, Dreiblat-

ELISE, Gaichycpatshosie cee 130
Bre, WANOWY ococecacc 5
PANemivy nestor ee 157
Pinette de prairie 185
Pitcher-plant ......... 192
Plant-of-peace ........ 42
RoplarWalldiaeeeeeee 77
Possession vine ...... 169
Powitch tree J........ 97
Prairie gourd ....... 180
IDK WER, Gonooudcc6 34
Prairie indigo ........ 105
Pride-of-Barbadoes .... 107
Queen’s fettle ....... 81
Queue de chat ....... 15
Queue de rat ........ 148
Queue de renard ..... 14
Ouillicane es ee Bik
Racine d’ours ........ 80
Ragweed, Short ...... 183

Raspberry, Boulder .... 93

Raspberry, False ..... 93
Rattlesnake weed ..... 193
Red beech ........... 57
Red coon-root ........ 82
Red death ........... 49
Red huckleberry ...... 161

Red mountain-dewberry 96

Red-ink berry ........ 68
Red-stemmed peavine.. 111
Redtop! ise eae 41
Reiss, Wilde ........ 29
RicesBlackj eerie 29
Riz de l’ane ......... 27
Riz farouche ......... 27
Riz sauvage ......... 27
Root, Beaver ........ 74
Root, Red coon ...... 82
Rose, Cotton ......... 133
Roseau, cane ........ 35
Rosemary, Sand Hill.. 123
Rothe Ceder ......... 13

Rothe Kalbsblume .... 83

Rothe Weide ........ 55
ounidenushpeeseeeeeoe 38
Rein, IE oooooo0cod 15
Rushy Roundy see eee 38
Riga, SHG. scosococxcc 45
Sac a commis ....... 158
Sacalhuistamemeernreiaee 33
Sage, Wood ......... 112
Sakakomigmeeneeierniers 158
Sakkakomi-Pflanze .... 158
Sand hill rosemary ... 123
Seite @ackye solocoosasoo 135
Salle UIA bolobosoadoo 45
Salle GRASS coococoooc 65
Salt=manshpaeernccicrerciea. 30
Sand myrtle ......... 190
Seicsy Jae corooaccoo 121
Sageloiee saccoscccce 101
Schachtelhalm ....... 3
Schlangenwurzel ...... 63
Schmalblattrige Weide 54
Schwarz wallnussbaum 52
SEF GEMS Goonoodsocon 30
Sea-fennel ........... 65
Shad-blossom ...... 86, 102
Short ragweed ....... 183
Slory, WE cosccouec 128
Silvery oleaster ...... 141
Sloe berry ........... 163
Small bonnet ........ 71
Small mountain-black

berry: .Shiveciaws Paacs 95

168

snotweed! = Fo u.5cs ccs 69 Tree, Beef-suet ....... 1AZD aWieed Snot seer 69
Snow-ball, Swamp .... 87 Tree, Boojum ........ 137. Weed, Stagger ....... 85
SOUt 2rdaSSs eee 23 Tree, Buck-wheat .... 127 Weed, Telegraph .... 188
Spalt. cake Aes hemos 60) -=irees -Powitchy 4-2 97 Weed, Tumble ....... 66
Spice; Swap) ----e- = 129) sErout lily, 2s -noeeieene 47 Weide, Rothe ........ 55
Spierstande, Wieden- fulpenbauml (coe oee 77 Weide, Schmal-

blatterige) 222.5 << 91 Tumbleweed ....:.-.- 66 blattriges Se cece 54
Splatter-dock) -s2-2 421 75. Wuarkshead' a... sick 40 Weideneiche ......... 58
Spoonwood eee eee eek 76 Turnip, Angel’s ...... 168 Weiden-blatterige Eiche 58
Squaw-berry ......... 126i LurnipsWaildmecsseeee 106.) Wermuth) 3---4> eee 196
Staggerweed ......... 85) BUWlimes Wahue eocceace 61. SWihipy 9 -3)45-co cere 45
Star bur*sersaoete seek 191” WWskotask 2222 scene 148 White beech ......... 57
Stechaplelly man aancere 73% Manilla) ss seaecte sec ne 198 White bull-tongue .... 20
Stinking willow ...... 141 Vetch, Augusta ...... 5s White: death) sense 51
Sloraxbaumy eee eee 88 Vine, Possession ..... 169 Whortle, Ground ..... 162
Sunshine bush ....... 165, Vanes View oo... ese 169" Wihorts) = oases cere 162
Swamp snow-ball ..... S7 = Wannellay eae eee 198 Wiedenblatterige
Swamp spice ........ 129 Wacholder, Kriechende 11 Spierstande ........ 91
Sweet) cedar Sscncanc JO! Wachsbaumpemaee ooo. 53) WaldWolive® < a-c5 scl 166
Tabackspflanze ....... 174 Waggina-gan ........ GaWaldgpoplars a+ eeeeoee 77
Tall blue huckleberry . 159 Wahu-Ulme.......... 6f Wald tarnip eee 106
Wap-palt sesso ee 141 Wallnussbaum, Schwarz 52 Wild winter-pea ...... 116
Meas ein diany ysis oleic 153. Wasser-Ahorn ........ 90 Wildcat grass .......- 32
Teddy-bear cholla .... 138 Wasser-Buche ....... 90 Wilde Reiss ......... 29
Telegraph weed ...... 188) Water beech jc cicic.cncr 57. Willow, Stinking .... 141
Texas doveweed ...... 120° Water berry) .<..-.<<0-- 179 Wintergreen, Mountain 156
Texas Spur’ syst cee 191 Water parsley ....... 146 Winter-pea, Wild ..... 116
Three-cornered grass... 40 Water-velvet ......... 2 Wire crass’ Se eee 32
Three-square, Leafy .. 40 Weed, Billy goat ..... OD SWioodisagessneseiereielce LUZ
Thimbleberry ......... 93°) Weed: ‘Button .22.- ..- 194 Wood, Spoon ........ 76
LUO. EBongoesaeoe 169 Weed, Camphor ..... 189 Woods clover ........ 108
iEra-ehiny .~\. =-.-asaces<s 46. Weed) Coffee’ -225---- I) Wie Eee So 5eacdosd0 119
Traubenkirsch ....... 100) wWieeds ebure see eee 186) Wiaparshiibeese eee 128
Mreadsatty peer 171 Weed, Mesquite ...... 104 Yellow grass ......... 26
Tree, Aniseed ........ 78 Weed, Moses ........ 34 Mellow) pine ssecee ee 5
Tree; ‘Bead! 2. ssh wesc 118 Weed, Rattlesnake .... 193 Zauberhaselnuss ..... 89

U. S. FisH AND WILDLIFE SERVICE
Cuicaco, ILLINOIS

Polypetalous Forms of Vaccinium

W. H. Camp anp C. L. GILty

During the course of the last decade one of the authors of this
brief paper has been giving some consideration to the Ericales. In
this study abnormalities of several types have been noted in various
groups. Among these is the polypetalous condition in Vaccinium.

In the genus Vaccinium the corollas are normally gamopetalous,
yet the polypetalous condition is closely approached in two groups: .
namely, the circumpolar subgenus O-xycoccus, and the subgenus
Oxycoccoides (= Hugeria Small), the latter found in southeast-
ern North America and eastern Asia. In these two groups the
corollas are not strictly polypetalous; instead, the corolla seg-
ments are deeply divided. It is to be noted that in Befaria, ap-
parently one of the more primitive of the living ericalean genera,
the corolla is always polypetalous and that this condition is cor-

169

related with an unstable number of parts of various of the floral
organs.’ Whether the deeply divided condition of the corolla in
the subgenera Oxycoccus and Oxycoccoides of Vaccinium is a
primitive character, or of more recent origin, is outside the present
discussion. We are here concerned solely with the phenomenon of
polypetaly in the subgenus Cyanococcus—the true blueberries—ot
eastern North America.

The usual gamopetalous corolla of Vaccinium indicates its der-
ivation from a polypetalous type by the marked apical lobing and
the folds which, in some species, lead from the sinuses toward the
base (figure 1g). It is therefore not surprising that, on occasion, the
normal gamopetalous corolla splits into its fundamentally component
parts. This situation was recorded in the literature a few years ago
by Weatherby. The description of this material indicates that the
polypetalous condition was not completely stable, for various types
of segmentation were present on the same plant.”

For the last several years the present authors have watched an
abnormal clone which grows naturally in the woodland of the New
York Botanical Garden on the hillside just south of the Arch
Bridge, and which in consecutive years has produced polypetalous
flowers (figures la-f). It is a low-growing form apparently derived
from Vaccinium torreyanum, which is common in the area.’

1 Camp, W. H. Studies in the Ericales. A discussion of the genus Befaria
in North America. Bull. Torrey Club 68:100-111. 1941. ;

2 Weatherby, C. A. A teratological form of Vaccinium pennsylvanicum.
Rhodora 29:237, 238. 1927. .

377. torreyanum is part of the complex which, in the manuals, has been
called V. vacillans. The “vacillans-complex,” spreading over much of eastern
North America, contains the following: the southern and central Appalachian
V. pallidum Ait. (not V. pallidum of the manuals), a somewhat coarse shrub
with yellowish branches, sometimes ascending to two or even three feet; the
more northern, northeasterly and Outer Piedmont V. torreyanum Camp with
its delicate, mostly greenish-barked branches rarely rising to more than
eighteen inches; the broad- and veiny-leafed ”. subcordatum (Small) Uphof,
a plant apparently confined to the Cumberland Plateau and several of its out-
liers; and ’. viride Ashe and ’. missouriense Ashe, both of which have their
primary centers somewhere in the Ozark Plateaus. These last two are dis-
tinguished from the others by their puberulent leaves, the coarser V. viride
apparently bearing much the same relation to . missouriense that V.
pallidum does to V. torreyanum. Whether it will be advisable in the future
to keep these as nomenclaturally separate species, or to recognize them as
parts of a widespread and regionally variable species, will be decided only

170

There is little need to give any detailed description of this plant
except to call attention to the fact that the five corolla segments of
each of the flowers examined were separate to the base. This con-
dition was obvious even in the bud (figure 1b). As in the case of
the plant noted by Weatherby (loc. cit.), the anthers apparently
were abortive (figures 1d, e). Whether the sterility extended to the
egg-apparatus has not been determined, although it is our observa-
tion that this plant does not set fruit. Furthermore, attempts to
produce fruit through the medium of artificial pollination have been
unsuccessful. However, this last is not a final conclusion, the at-
tempts so far having been attended by conditions which admittedly
were not ideal. In brief, we are not as yet convinced that this clone
is incapable of setting fruit.

In addition to the polypetalous condition, one other anomalies
should be noted. In sectioning the hypanthium of a series of the
flowers of this clone it was found that the ovary of each had but
four carpels (figure 1f), instead of the five carpels normal for /.
torreyanum and its close relatives (figure 1j). This, however, is by
no means unusual in the genus Vaccinium. It is quite common in
certain species and is, in fact, a standard character of others.
Nevertheless, this does indicate that the disturbance resulting in the
polypetalous condition can also influence the number of carpels in
the ovary. In this connection, it is of interest to note that various
of the nearly polypetalous members of the subgenera Oxycoccus
and Oxycoccoides are tetramerous, with the pentamerous condition
being the abnormal form. Whether this condition is merely coin-

after further and much needed cytological studies of these entities have been
made throughout their entire distributions. In addition to the foregoing, the
“Yacillans-complex” contains V. tallapusae (Cov.) Uphof, a derived tetraploid
of the southern Appalachians which is best developed in Georgia; V. alto-
montanum Ashe of the southern Appalachians may also be a derivative of
this group. V. vacillans var. crinitum Fernald, with which V. missouriense
and V’. viride have been confused, appears to be a series of hybrids and
ecologically selected segregates from crosses between the markedly different
V. torreyanum and V. atrococcum, both of which are diploid (n— 12) and
known to be interfertile. The “high-bush” diploid V. atrococcum (A.Gr.)
Heller apparently does not enter the ranges of V. missouriense and V. viride,
being primarily an east-coast species; westward, it has been confused with
V. arkansanum Ashe, and with the “arkansanoid’” members of the tetra-
ploid V. corymbosum L.

171

aS)

er Z
SSS = -
SS,

eed

GL.Gil\
— 1942.

Ficures la-f: Material from a clone of Vaccinium torreyanum growing
naturally in the woodland of The New York Botanical Garden which, over a
period of years, has borne polypetalous corollas. Ficure la: Habit sketch
of one branch, natural size. Ficure 1b: Indicating the position of the petals
in the bud, X4. Ficure 1c: The fully opened flower, X4. Ficures ld, e:
Two views of a stamen showing the abortive anther, X 8. Ficure 1f: Diagram-
matic cross-section of the hypanthium showing the four-carpeled ovary.
Ficures lg-j: The flower from a normal clone of V. torreyanum growing near
the former. Ficure lg: External view of the flower at anthesis, X4. FIGURE
1h, i: Two views of a stamen showing the normal anther, X8. Ficure 1j:
Diagrammatic cross-section of the hypanthium showing the normal five-
carpeled ovary. As is common in Vaccinium, the “false partitions,’ coupled
with the elongated placentae, give the appearance of a ten-carpeled ovary;
the vascular structure (not included in the diagram) indicates its true nature.

172

cidental, or of fundamental evolutionary significance, is not known.

In addition to the clone here under consideration, one of the
authors of this note has found much the same condition in other
species. A collection of Y. atrococcum from central New Jersey has
been seen where the corollas were still gamopetalous, but with the
segments so poorly united that even a slight pressure would cause
them to fall apart. It was also found in a clone of lV’. brittonw Porter
on High Point in the Kittatinny Mountains of extreme northern
New Jersey. In lV. brittoni the condition was variable, much as in
the material mentioned in Weatherby’s discussion. Incipient poly-
petaly has also been observed in other species of the genus but
seldom in so complete a condition as the material figured in this
paper.

It is therefore obvious that the individual plant in the genus
Vaccinium, through some disturbance, may produce polypetalous
corollas. The genetics of the situation so far has not been studied, for
anther deficiencies often accompany the condition. There is also
some slight but not conclusive evidence that the plants may also be
sterile to viable pollen.

The nomenclature of such forms should be considered. Similar
plants with at least deeply divided gamopetalous corollas have been
the basis of such entities as Rhododendron linearifolium Sieb. &
Zuce. (in which there is also some disturbance of the leaf form),
Kalnua latifolia var. polypetala Nichols, and Rhododendron at-
lanticum forma tomolobum Fernald. There is evidence that the
precise application of these epithets requires that they be used to
refer only to single clones. Since this is the case—and essentially
the same manifestation is the basis of a species, a variety and a form
—it would seem only logical that some other category be selected to
designate the polypetalous condition in the genus Vaccinium, and
thus complete the nomenclatural cycle.

The foregoing is said less in jest than may at first appear to be |
the case. It is not the primary purpose of this paper to discuss the
proper nomenclatural disposition of such obviously aberrant ma-
terial. However, it would seem that nomenclature should at least
be functional ; that its purpose is not only the listing of differentiable
entities, but also that it should in some way indicate their proper
relationships. Therefore, it is our opinion that, where there is need,
an organism should have a name but that the category to which it

173

is assigned should have some biological significance in a system of
nomenclature. This is equally true of horticultural material and of
organisms growing naturally under feral conditions. In this in-
stance, it is doubted whether a single aberrant clone—as Rhododen-
dron linearifolium—deserves specific rank, particularly when the
normal form has to take nomenclaturally subsidiary rank under it
as a variety.”

Were the polypetalous individuals of Vaccinium to be brought
into cultivation—and if propagated by asexual means—they would
deserve no more than the category of “lusus” as originally defined
by DeCandolle. Yet it is admitted that this material is of little or no
importance either as a horticulturally or otherwise useful plant-type.
It is therefore thought best not to further encumber the literature
of the group with a series of subspecific names which, for the pres-
ent, would seem to serve no practical purpose. The polypetalous
condition in Vaccinium is perhaps of some interest from the botan-
ical standpoint and it is for this reason that this paper has been
prepared. Further study of the phenomenon may lead to other work
on the origin of somewhat similar forms and thus perhaps shed
light on one phase of the general evolution of the group. Some future
worker dealing with these matters may find it desirable to give
names to such individuals, if only to particularize and expedite his
discussions ; for the present—to us at least—they are only items
of general biological interest and therefore scarcely worthy of
nomenclatural recognition.

THe New York BotANiIcaAL GARDEN
New York, NEw York

*#Rehder (Man. Cult. Trees and Shrubs, 1940) begins the description of
Rhododendron linearifolium as follows: “A garden form of the following

.” The following entity is R. linearifolium var. macrosepalum (Maxim.)
Mak. One wonders how the apparently basic, normal material can be con-
sidered a variation of an obviously derived and abnormal, vegetatively
propagated clone (and therefore, biologically, an individual) except where
nomenclature is an end in itself rather than a means by which information
can be better organized. The writers of this note bow to the accusation that
they hold to the principle that nomenclature, as such, should be a tool in the
science of systematics, rather than the view that systematics is a mental
diversion appended to the science of nomenclature.

Carex aestivalis and Carex lurida var. gracilis on the
Glaciated Allegheny Plateau

Rospert T. CLAUSEN

Alma Hill, in Allegany County, is one of the highest hills
(elev. 775 m.) in western New York. The flora and fauna both are
strongly characteristic of the Canadian Life Zone. Birds such as the
Olive-backed Thrush, Winter Wren, Blue-headed Vireo, and Junco
seem to be common breeding species there. Lycopodium annotinum
var. integrifolium, Dryopteris Phegopteris, Schizachne purpuras-
cens, and Milium effusum further suggest the northern character
of the region. On the wooded slopes, Carex radiata, a species which
is rare in central New York, is frequent. Carex aestivalis, even rarer
in central New York, occurs in dry rocky woods near the western
base of the hill. Along a brook, also on the west side of the hill,
at an elevation of 580 m., Carex lurida var. gracilis occurs.

Data at hand indicate that both Carex aestivalis and C. lurida
var. gracilis are infrequent and local on the Glaciated Allegheny
Plateau. In the herbarium of Cornell University, neither sedge is
represented from this plateau in Pennsylvania or Ohio. Mackenzie
(1931-35) did not mention having seen specimens of either from
Ohio, although it is possible that both may eventually be discovered
in the northeastern part of that state.

House (1924) recorded the range of C. aestivalis in New York
as “Dutchess county, the Catskill mountains and Otsego county,
southward.” The two localities in Otsego County, Worcester and
East Worcester, are both on the Glaciated Allegheny Plateau. To
these may be added the following localities from which specimens
are available in the herbarium of Cornell University: woods,
4 Town Schoolhouse, Sempronius, 3 miles east of Moravia, Cayuga
County, July 11, 1882, herbarium Charles Atwood; Dresserville
Gulf, town of Sempronius, Cayuga County, September 12, 1896,
herbarium Atwood; dry steep, shaded, sandy-clay bank, ““The Nar-
rows’ Slaterville to Caroline Center, Caroline, Tompkins County,
July 13, 1919, A. J. Eames, K. M. Wiegand, & L. F. Randolph
11594 ; roadside slope, wooded ravine just east of Bald Hill, Caro-
line, Tompkins County, July 2, 1936, M. W. Allen 19329; dry
rocky woods near base of western slope of Alma Hill, Allegany

174

N75)

County, June 18, 1939, R. T. C. 3917; and wooded bluff by creek,
Sinclairville, June 24, 1924, K. M. Wiegand 15256. A further
locality, reported by Zenkert (1934), is South Wales in Erie
County. Hamburg, also cited by Zenkert, is on the Great Lakes
Plain. From just south of the terminal moraine, in the Allegany
State Park, House and Alexander (1927) reported C. aestivalis as
frequent.

House (1924) reported Carex Baileyi (C. lurida var. gracilis)
southward to Greene and Herkimer Counties, also from Campville,
Tioga County. House and Alexander (1927) reported this same
variety as common in the Allegany State Park, an unglaciated
area. On the Glaciated Allegheny Plateau, where typical Carex
lurida is common, the var. gracilis seems to be rare. Records are
available only from Allegany, Chemung, and Tioga Counties. The
following two collections are in the herbarium of Cornell Univer-
sity: open swaly clearing in white oak woods, Comfort Hill,
Chemung County, June 29, 1938, S. J. Smith & Harvey Scudder
933; and woods along brook on west side of Alma Hill, Allegany
County, June 18, 1939, Rk. T. C. 3920. In this herbarium there are
several specimens intermediate between Carex lurida var. typica
and var. gracilis. These are from Rutland County, Vermont; Nor-
folk County, Massachusetts ; Hartford County, Connecticut ; Albany
and Fulton Counties, New York; and Haywood County, North
Carolina. These support Wahl’s (1940) statement that “Carex
Baileyi (C. lurida var. gracilis [Boott] Bailey) is very closely
related to C. lurida.’” In counting chromosomes of Carex lurida,
Wahl found haploid numbers of 32 and 33 for three plants of the
typical variety. A plant of var. gracilis was n = 34. Wahl’s data
for various Carices reveal that plants which are morphologically
similar and which certainly belong to the same taxonomic species
may differ in having one or two chromosomes more or less. Accord-
ingly, the number 34, mentioned above, does not strengthen the
case for treating var. gracilis as a species, since typical C. lurida
already is known to be either n = 32 or 33. Though found primarily
in the northern part of the range of the species and usually at
higher altitudes, the var. gracilis can not be regarded as a strongly
geographical entity, since its distribution lies entirely within the
area of typical C. lurida.

176

Literature Cited

House, H. D. 1924. Annotated list of the ferns and flowering plants of New
York State. N. Y. State Museum Bull. 254. p. 1-759.

House, H. D. and Alexander, W. P. 1927. Flora of the Allegany State Park
Region. New York State Museum Handbook 2. p. 1-225. illus.

Mackenzie, K. K. 1931-35. Cyperaceae-Cariceae. N. Am. Flora 18: 1-478.

Wahl, Herbert A. 1940. Chromosome numbers and meiosis in the genus Carex.
Am. Jour. Bot. 27: 458-470. t. 1-2.

Zenkert, C. A. 1934. The flora of the Niagara Frontier Region. Bull. Buffalo
Soc. Nat. Sci. 16: 1-328.

CoRNELL UNIVERSITY
IBEUNOR, IN, 3%,

Papers on the Flora of Alaska—I.
The Genus Cicuta

J. P. ANDERSON

The species of Cicuta are of great importance on account of
their very poisonous properties. Losses of cattle directly attributed
to poisoning by C. douglasii (DC.) Coult. & Rose have occurred
in southeastern Alaska. There have been rumors of losses elsewhere.

In a recent study, Mathias and Constance (1) have reduced
the American species of the genus to seven. Of these, three occur
in Alaska. The following key covers these three species :

Fruit longer than wide, leaflets 2-4 times as long as wide.

C. maculata

Fruit shorter than wide, leaflets 5-10 times as long as wide.

& mackensieana

Fruit about equal in length and width, leaflets 134-2%4 times as long
as wide. C. douglasu

This is probably the first report of C. maculata L. from Alaska,
although C. virosa L. reported by Porsild (2) from Hot Springs
on the Tanana River undoubtedly was this species. The author
first collected it at Knik on Knik Arm of Cook Inlet in 1931 (1382).
In 1935 it was collected at Circle Hot Springs (2616), as again in
1941 (7560). A collection at Hyder in 1939 (5501) is rather im-
mature but seems to be this species. A visit to Manly Hot Springs
(also known as Tanana Hot Springs) in 1941 revealed its presence

177

there (7075). It is to be noted that the most northerly stations
are at hot springs where other more southerly species of plants
also occur.

Cicuta mackenzieana Raup is the most widely distributed mem-
ber of the genus in the territory. The author first collected it at
Matanuska in 1922. The following collections have since been made:
Matanuska (1103); College, near Fairbanks (1258) ; Circle, on
Yukon River (2595); mile 312, Richardson Highway (2686) ;
Willow Creek, mile 92, Richardson Highway; Valdez (2588) ;.
Unalakleet (5106) ; Eklutna (6940) ; Takotna (7352) ; Talkeetna
(7580). An immature specimen collected at Hope (6695) seems
to belong here.

a DRO e

a. Leaflet of C. douglasti. b and c. Leaflets of C. mackenzieana. d. Leaflet
of C. maculata. All drawn by Dr. Ada Hayden.

Cicuta douglas (DC.) Coult. & Rose seems to be confined to
southeastern Alaska. Collections were made at Lemon Creek (784)
and at Mendenhall (783), both near Juneau ; Haines (1570) ; Skag-
way (1733) ; and Echo Cove, Lynn Canal (6034).

All specimens cited were collected by the author and are in
his herbarium now deposited in Iowa State College at Ames.

178

So far as their occurrence in Alaska is concerned, the species
are very distinct and can be readily separated by vegetative char-
acters alone. Cicuta douglasu and Cicuta maculata grow up to
2 meters tall and Cicuta mackenzieana up to 1% meters. All have
bipinnate leaves which often appear to be ternate-pinnate. In
C. douglasii the leaflets are lanceolate, 1-5 cm. wide X 2Y%-9 cm.
long, serrate to doubly serrate or even incised and with sharp
teeth. In C. maculata the leaflets are more narrowly lanceolate,
.8-3 cm. X 3-9 cm., rather evenly serrate with sharp, mostly out-
ward pointed teeth. The leaflets of C. mackenzieana are narrowly
lanceolate to linear, .2-2 cm. & 2-10 cm., with rather remote,
sharp, forward-pointing teeth, these being rather small on the
more narrow leaflets. Fruits of C. douglas are deeply grooved,
about 234 mm. long and wide. C. mackenzieana also has deeply
grooved fruits which are about 2% mm. wide < 2 mm. long. In
C. maculata the fruit is not grooved, the space being filled by thick
corky ribs. The fruit measures about 234 mm. wide X 3% mm.
long. All the species are found growing in shallow water, some-

times ascending into mud, but never in well drained situations.
This habitat is quite different from that of other tall growing
members of the same family, which are always found on better
drained soils.

Literature Cited

Mathias, Mildred E. & Lincoln Constance. 1942. A synopsis of the Ameri-
can species of Cicuta. Madrono 6: 145-151. Pl. 14.
Porsild, A. E. 1939. Contributions to the flora of Alaska. Rhodora 41: 266.

Botany DEPARTMENT
Iowa STATE COLLEGE
Ames, IowA

Some New Forms from the Middle West

NorMAN C. Fassett

NapaeEa pioIca L., f. stellata, n.f., foliorum setis stellatis, ramis
0.2-0.5 mm. longis, rare simplicis 1.0 mm. longis——Along a rail-
road 3.8 miles west of Cross Plains, Wisconsin, August 16, 1942,
N.C. Fassett, no. 22057 (Type in Herb. Univ. of Wis.).

N. dioica occurs with two quite distinct types of pubescence. In
some plants the lower leaf-surfaces have straight appressed simple
hairs a millimeter long, with only occasionally a stellate trichome.
In others these simple hairs are nearly or quite lacking except on the
larger veins, and are replaced by close stellate hairs with short
branches. The first type is represented in the Herbarium of the
New York Botanical Garden by a sheet from Pennsylvania, one
each from Cincinnati and Peoria, and by two cultivated plants.
The second is represented by collections from Ohio, Indiana, IIli-
nois, Wisconsin and Iowa. In the Herbarium of the University of
Wisconsin there are from this state 3 sheets with simple pubescence,
and 17 with stellate trichomes predominating. It is therefore evi-
dent that both forms occur in the Middle West (where, despite

Leaf of Napaea dioica, X\4.

179

180

the statement in Gray’s Manual, the plant is by no means rare).
Linnaeus did not mention the type of pubescence, and most subse-
quent authors merely specify “scabrous” or “roughish.” Sprengel,
Syst. Veg. 3:122, describes Sida dioica as “S. herbacea hirsuta”
perhaps implying simple hairs. Since the simple-haired plant is
certainly present in the east it is taken as the typical form, and the
other here described as f. stellata.

Two Mass Collections have been made, with upper and lower
leaves from a single plant in each clone. One from Cross Plains,
Wisconsin, consists of 12 pistillate plants and 2 staminate plants ;
the other, from near Black Earth, shows 8 pistillate and 3 staminate
plants. These 25 individuals are all f. stellata, but this observation
does not imply that the two forms may not grow together in some
regions.

The figure in Britton & Brown’s illustrated Flora purporting
to illustrate Napaea will look strange to anybody who is familiar
with the deeply 7-lobed leaf of that plant. Perusal of the material in
the New York Botanical Garden brought to light a specimen which
closely matched the drawing, and was obviously the original ; it is
Sida hermaphroditica. Since there seems to be no readily available
illustration of the very characteristic leaf of Napaea dioica, one is
here presented.

ASARUM CANADENSE L., var. ACUMINATUM Ashe, f. Prattii, n.tf.,
calycibus viridibus non purpureis—Wooded bank, Green Lake,
Wisconsin, May 20, 1938, C. H. Pratt & N. C. Fassett, no. 22001
(Type in Herb. Univ. of Wis.). This clone of Wild Ginger with
green flowers has been under observation by Mr. Pratt for several
years. It seems to be quite analagous to A. caudatum f. chloroleucum
Palmer in St. John, Proc. Biol. Soc. Wash. 41:193. 1928. -

LaTHyRUS JAPONICUS Willd., var. GLABER (Ser.) Fernald, f.
spectabilis n.f., corollis coccineis—Cobblestone beach of Lake
Superior, 12 miles east of Grand Marais, Minnesota, July 12, 1938,
N. C. Fassett & J. T. Curtis, no. 22000 (Type in Herb. Univ. of
Wis.). The deep crimson flowers of this plant were conspicuous
among the ordinary purple-flowered individuals of Beach Pea.
When pressed, they became a very deep blue.

ZANTHOXYLUM AMERICANUM Mill., f. impuniens, n.f., ramulis
inermis.—Three miles north of Wisconsin Dells, Juneau County,

181

Wisconsin, September 13, 1938, N. C. Fassett G J. W. Thomson,
Vir, WO, ALBZZ (Anes, ta leleno. © Wi), (Oh wats SS Saesis Oi
Prickly Ash from Wisconsin. 4 lack the sharp prickles usually char-
acteristic of this shrub.

MIMULUS RINGENS L., f. roseus, n.f., corollis roseis.—Sandy
shore of the St. Croix River, Evergreen, St. Croix County, Wiscon-
sin, July 31, 1934, N. C. Fassett, no. 21821 (Type in Herb. Univ.
of Wis.).

DEPARTMENT OF BOTANY
UNIVERSITY OF WISCONSIN

Clarence J. Elting and his Herbarium

Homer D. House

The New York State Museum has recently acquired as a gift
from Mrs. Elting, the herbarium of the late Clarence J. Elting of
Highland, Ulster County, New York. The collection is noteworthy
among small local herbaria because of the careful preparation,
preservation and correct identification of the material according to
the current floras of his day. Most of the specimens exhibit both
flowering and fruiting specimens, pressed and mounted with a skill
rarely seen in such collections, with fairly accurate data as to
locality and date. Over 90 percent of the specimens were collected
in eastern Ulster County, New York. The remainder are from
Mohonk, Minnewaska and Denning in Ulster County, with a few
from across the Hudson River in Dutchess County. Important as
a contribution to the local flora of New York City and vicinity it
calls for some mention of the principal items among the 1075 speci-
mens.

Clarence J. Elting was born October 13, 1860, at Highland,
Ulster County, New York, where he spent most of his life until
his death on May 28, 1942. His interests were mainly amateur
photography, botany, genealogy and local history. He was a member
of the local historical societies and well known locally as an authority
upon such matters. The New Paltz Independent of Thursday,
June 4, 1942, contains additional information regarding his life
and activities.

182

Some details are necessary in connection with the localities
where most of his plants were collected. Bailey’s Gap is the highest
point on Route 55, between Highland and Clintondale, and is
3 miles west of the Hudson River. Saxton’s Pond is in the town-
ship of Lloyd and is now designated on topographic maps as “Lily
Lake.” Pine Hole is an extensive swamp and bog area on the head-
waters of the Swartekill about 2 miles south of Ohioville, a hamlet
about 2 miles east of New Paltz. Butterville is a hamlet about 2
miles west of New Paltz. Dashville Falls is near Dashville on the
Wallkill River. Black Pond is in the northern part of Lloyd town-
ship. Bull Run is in the township of Denning, about 3 miles north
of the Sullivan County line. Claryville, Libertyville, Lloyd, Mohonk,
Minnewaska, New Paltz, Clintondale, Marlboro and Milton are
all in Ulster County and easily located on most maps. Fallsburgh
is in northern Sullivan County and Millburn is a hamlet in Wallkill
township, Orange County.

Most of the specimens are numbered apparently according to
their sequence in his Gray’s Manual. The herbarium was started
in 1892 and active collection was continued until 1903, with fewer
collections in 1907 and some scattered ones as late as 1923 and 1925.

The following species are selected from Mr. Elting’s herbarium
as most worthy of permanent record:

ANCHISTEA VIRGINICA (L.) Presl. Swamp near Mohonk, 3584, August 10,
1896

ASPLENIUM MONTANUM Willd. Near Mohonk, 3540, September 5, 1901

PoLysTICHIUM BRAUNI (Spenner) Fee. Mountain woods near Shokan, 3521,
May 15, 1903

WoobsIa oBTuSA (Spreng.) Torr. Rocky woods, Highland, 3580, July 8, 1893

BotTRYCHIUM LANCEOLATUM (Gmel.) Angstr., var. ANGUSTISEGMENTUM
Pease & Moore. Woods near Highland, August 7, 1900

BoTtRYCHIUM MULTIFIDUM (Gmel.) Rupr., var. SILAIFOLIUM (Presl.) Broun
(Ind. N. Am. Ferns 41, 1938). Woods near Highland, 3592, August 29,
1898

PICEA MARIANA (Mill.) B. S. P. “Pine Hole’ swamp, 2 miles south of
Ohioville, 2515, May 20, 1897

SPARGANIUM AMERICANUM Nutt. Shallow water near Highland, 2772, July 10,
1893

POTAMOGETON PECTINATUS L. Shallow water, Libertyville, 2851, July 24, 1898

SAGITTARIA SUBULATA (L.) Buchenau. Borders of Hudson River near High-
land, 2808, August 20, 1892

183

SAGITTARIA LATIFOLIA Willd., f. DIVERSIFOLIA (Engelm.) Robinson. Hudson
River near Highland, 2810, August 8, 1892

ANACHARIS OCCIDENTALIS (Pursh) Marie-Victorin. Tidal marsh on Hudson
River near Highland, 2531, July 28, 1894

Pantcum AsHEI Pearson. Woods near Highland, 3411, June 19, 1901

MUHLENBERGIA RACEMOSA (Michx.) B. S. P. Borders of Saxton’s Pond,
Lloyd, 3395, September 14, 1896

CALAMAGROSTIS CINNoIwES (Muhl.) Barton. Minnewaska, 3302, August 25,
1896

ELEUSINE INDICA Gaertn. Waste ground, Highland, 3328, August 13, 1895

TriopIA FLAVA (L.) Hitchc. Near Esopus, 3493, October 10, 1899

GLYCERIA FLUITANS (L.) R. Br. Shallow water near Highland, 3364, June 8,
1899. Well marked by the large spikelets and long lemmas.

Bromus HorDACEUS L. Roadside near Highland, July 10, 1907

ELtymus Wrecanpit Fernald (Rhodora 35: 192, 1933). River shore near
Highland, 3329, July 29, 1894

‘Hystrrx pATULA Moench. Woods near Highland, 3274, July 12, 1894

ELEOCHARIS CAPITATA (L.) R. Br. (E. tenuis of N. Y. Reports). Wet soil,
Highland, 3144, June 24, 1896

STENOPHYLLUS CAPILLARIS (L.) Britt. Dry soil, Highland, 3170, August 21,
1896

ScirPUS HUDSONIANUS (Michx.) Fernald (Erioprorum alpinum L.) “Pine
Hole’ swamp, 2 miles south of Ohioville, 3159, June 3, 1896

ERIOPHORUM VIRIDI-CARINATUM (Engelm.) Fernald. “Pine Hole’ swamp,
2 miles south of Ohioville, 3164, June 3, 1896

RyNcHOoSPORA ALBA (L.) Vahl. Borders of Saxton’s Pond, Lloyd, 3180,
August 16, 1895

Mariscus MArRIScOIDES (Muhl.) Kuntze. Saxton’s Pond, Lloyd, 3102,
August 6, 1895

CAREX EXILIS Dewey. “Pine Hole” swamp, 2 miles south of Ohioville, 2924,
May 20, 1897

CAREX CEPHALANTHA (Bailey) Bicknell. Borders of Saxton’s Pond, Lloyd,
2921, June 5, 1897

CAREX TORTA Boott. Along stream, Bull Run, Denning, 3065, May 15, 1903

Carex Davistr Schw. & Torr. Meadow, Hackensack road, southeast of
Poughkeepsie, Dutchess County, 2908, June 1, 1899

CAREX PLANTAGINEA Lam. Woods near Bull Run, Denning, 3004, May 12,
1903

CAREX GRISEA Wahl. Moist woods, Libertyville, 2967, June 17, 1899

CAREX CRYPTOLEPIS Mackenzie. Wet soil near Highland, 2930, June 14, 1899

CAREX LASIOCARPA Ehrh. Borders of Saxton’s Pond, Lloyd, 2927, June 5,
1897

CAREX SQUARROSA L. Woods near Highland, 3078, August 29, 1905

Carex Grayit Carey. Wet woods, New Paltz, 2946, June 23, 1902

ARISAEMA PUSILLUM (Peck) Nash. Wet woods, Highland, 2779, July 10,
1893

184

Peck (N. Y. State Mus. Bul. 67: 20, 1903) reports specimens con-
tributed to the State Herbarium by Mr. Elting, collected in June, 1902.
The type was collected at Millbrook, Dutchess County, by Fred. Thorne
of New Paltz (51st Ann. Rep’t N. Y. State Mus. 275, 297), but the
herbarium specimen is mistakenly labelled as having been collected by
“FE. Thomas.” Other New York collections in the State Herbarium are:
Bedford Park, Bronx, Nash, May 26, 1899; Sandlake, Rensselaer County,
Peck; Hewlett, Nassau County, Taylor (Torreya 9: 260, 1909).

ARISAEMA DracontiuM L. Wet soil, Libertyville, 2778, May 31, 1898

ORONTIUM AQUATICUM L. Saxton’s Pond, Lloyd, 2781, May 31, 1894

ERIOCAULON SEPTANGULARE With. Saxton’s Pond, Lloyd, 2870, July 5, 1895

Xyris FLExuOSA Muhl. Borders of Saxton’s Pond, Lloyd, 2709, August 6,
1895

CoMMELINA COMMUNIS L. Waste places, Highland, 2712, August 22, 1905

ZOSTERELLA DUBIA (Jacq.) Small. Shallow water, Libertyville, 2702, July 23,
1896

CyYPRIPEDIUM REGINAE Walt. Swamp near Highland, 2548, June 11, 1893

HABENARIA BRACTEATA ( Willd.) R. Br. Woods near Highland, 2555, May 22,
1893

HABENARIA FLAVA (L.) Gray. Woods near Highland, 2572, July 1, 1893

HABENARIA BLEPHARIGLOTTIS ( Willd.) Torr. Minnewaska, 2553, July 28, 1896

ARETHUSA BULBOSA L. Borders of Saxton’s Pond, Lloyd, 2536, May 13, 1896

PoGONIA OPHIOGLossompES (L.) Ker. Marsh near Highland, 2585, June 28,
1893

LIMODORUM TUBEROSUM L. Borders of Saxton’s Pond, Lloyd, 2537, June 20,
1895

SPIRANTHES PLANTAGINEA (Raf.) Torr. (S. lucida Ames). Wet soil,
Lloyd, 2590, June 13, 1901

GooDYERA PUBESCENS (Willd.) R. Br. Woods near Highland, 2551, July 17,
1894

CoRALLORRHIZA ODONTORHIZA Nutt. Woods near Highland, 2541, September 3,
1892

LIPARIS LILLIFOLIA (L.) L. C. Rich. Woods near Highland, 2574, June 6, 1903

APLECTRUM HYEMALE (Muhl.) Torr. Woods near Highland, 2535, May 26,
1895

MyricA PENNSYLVANICA Loisleur (J/. carolinensis of N. Y. Reports, see
Fernald, Rhodora 37: 423, 1935; 40: 410, 1938). “Pine Hole” swamp,
2 miles south of Ohioville, 2416, June 8, 1897

Carya ALBA (L.) K. Koch. Woods near Highland, 2412, May 24, 1893

CoRYLUS CORNUTA Marsh. Thickets near Highland, 2435, April 6 and June 20,
1893

BeETULA NIGRA L. Along stream near Libertyville, 2426, April 15 and July 10,
1893

< Quercus Scuvuetter Trelease (Q. bicolor X macrocarpa). Woods near
Highland, August 20, 1905

Morus ruBRA L. Woods near Highland, 2393, May 14, 1894

PARIETARIA PENNSYLVANICA Muhl. Dashville Falls, 2394, October 12, 1901

ARISTOLOCHIA SERPENTARIA L. Moist woods, Highland, 2317, August 10, 1895

185

PoLYGONUM TENUE Michx. Rocky slopes, Highland, 2298, September 17, 1896

PARONYCHIA FASTIGIATA Fernald, Rhodora 38: 421, 1936 (Anychia polygo-
noides Raf.). Dry soil near Highland, 2210, August 4, 1902

PARONYCHIA CANADENSIS (L.) Wood (Anychia dichotoma Michx.). Dry
woods, Highland, 2209, September 9, 1892

ARENARIA LATERIFLORA L. Woods near Libertyville, 261, June 28, 1897

ARENARIA GROENLANDICA (Retz.) Spreng., var. GLABRA (Michx.) Fernald.
Ledges near Minnewaska, 260, July 21, 1895

STELLARIA BOREALIS Bigel. Wet places in mountain woods near Claryville,
310, June 6, 1898

CERASTIUM NUTANS Raf. Moist places near Highland, 276, May 17, 1895

CERASTIUM OBLONGIFOLIUM Torr. Rocky soil, Clintondale, 274, May 18, 1898

LYCHNIS CHALCEDONICA L. Adventive, Highland, 284, July 1, 1894

AGROSTEMMA GitTHaAGO L. Field near Highland, 288, July 15, 1893

SItENE ArmertIA L. Adventive, Highland, 299, July 10, 1895

SILENE STELLATA (L.) Ait. f. Woods near Libertyville, 307, August 6, 1897

SAPONARIA VACCARIA L. Adventive, Highland, 296, August 26, 1897

DIANTHUS DELTOIDES L. Roadside near Highland, 280, June 10, 1893

CLAYTONIA CAROLINIANA Michx. Woods near Claryville, 320, May 12, 1897

NYMPHAEA opORATA Dryand. Black Pond near Lloyd, 108, July 11, 1893

RANUNCULUS FLABELLARIS Raf. Marsh near Highland, 65, May 14, 1894

RANUNCULUS AMBIGENS S. Wats. (R. obtusiusculus Raf. ?). Wet soil
near Highland, July 10, 1907

RANUNCULUS MICRANTHUS Nutt. Woods near Highland, 49, May 10, 1893

CLEMATIS VERTICILLATA DC. Rocky woods west of Highland, May 10, 1904

CopTIS TRIFOLIA (L.) Salisb. “Pine Hole” swamp, 2 miles south of Ohioville,
34, May 6, 1897

TROLLIUS LAXUS Salisb. “Bailey’s Gap,” near Highland, 82, April 21, 1903

DeLPpHINIUM Ajacis L. Garden adventive, Highland, 35, August 2, 1893

CIMICIFUGA RACEMOSA L. Woods near Libertyville, 24, July 17, 1897

DICENTRA CANADENSIS (Goldie) Walp. Mountain woods, Claryville, 129,
May 12, 1897

ADLUMIA FUNGOSA (Ait.) Greene. Rocky woods, West Mountain near High-
land, 122, July 14, 1895

BERTEROA INCANA (L.) DC. Roadside near Highland, 134, November 6, 1893

LUNARIA ANNUA L. Persistent, Highland, 189, May 8, 1898

ARABIS CANADENSIS L. Woods near Highland, 135, May 23, and August 10,
1894

POLANISIA GRAVEOLENS Raf. Along Hudson River near Highland, 215,
August 22, 1896

SARRACENIA PURPUREA L. Swamp near Highland, 112, June 6, 1893

DROSERA ROTUNDIFOLIA L. Borders of Saxton’s Pond, Lloyd, 830, July 8, 1895

DROSERA INTERMEDIA Hayne. Border of Saxton’s Pond, Lloyd, 828, July 8,
1895

SAXIFRAGA PENNSYLVANICA L. Swampy meadow near Highland, 809, May 19,
1896

HEUCHERA AMERICANA L. Thickets near Libertyville, 776, June 28, 1897

186

PARNASSIA AMERICANA Muhl. (P. caroliniana of N. Y. Reports). Wet soil,
Hackensack road, southeast of Poughkeepsie, Dutchess County, 787,
August 11, 1896

RIBES ODORATUM Wendl. Escaped and spreading, Highland, 793, May 15, 1893

SorBUS AMERICANA Marsh. Woods near Bull Run, Denning, 726, May 15, 1903

WALDSTEINIA FRAGARIOIDES (L.) Tratt. Woods near Fallsburgh, 767, May 15,
1897

PoTrENTILLA PALUSTRIS (L.) Scop. Border of Black Pond, Lloyd, 700, July 10,
1893

PoTENTILLA ANSERINA L. Along Hudson River, 2 miles north of Highland,
691, September 28, 1898

GEUM MACROPHYLLUM Willd. Mountain woods, Bull Run, Denning, 681,
May 12, 1903

GEUM RIVALE L. “Bailey’s Gap,” near Highland, 683, May 31, 1905

DALIBARDA REPENS L. Woods near Liberty, Sullivan County, 675, July 21,
1904

SANGUISORBA MINOR Scop. (Poterium Sanguisorba L.) Waste ground near
Marlboro, 711, October 14, 1903

SANGUISORBA CANADENSIS L. Wet soil, Lloyd, 710, August 18, 1903

CASSIA MARILANDICA L. Thickets near Libertyville, 517, July 15, 1896

CASSIA NICTITANS L. Field near Highlands, August 18, 1907

CROTALARIA SAGITTALIS L. Field near Highland, August 20, 1907

CoRONILLA VARIA L. Meadow near Highland, 525, August 12, 1892

DESMODIUM CANESCENS (L.) DC. Woods near Libertyville, 536, August 13,
1899

DESMODIUM BRACTEOSUM (Michx.) DC. Thickets near Highland, 539,
August 13, 1895

DESMODIUM LAEVIGATUM (Nutt.) DC. Woods near Highland, 540, August 13,
1895

DESMODIUM MARILANDICUM (L.) DC. Dry woods, Highland, 545, August 28,
1892

DeEsMopIUM RIGIDUM (Ell.) DC. Dry woods, Highland, 550, August 21, 1896

LESPEDEZA PROCUMBENS Michx. Dry woods near Highland, 579, September 11,
1896

LESPEDEZA REPENS (L.) Bart. Dry woods, Highland, 578, September 7, 1896

LESPEDEZA VIOLACEA (L.) Pers. Dry woods, Highland, 580, August 18, 1892

LESPEDEZA VIRGINICA (L.) Britt. Dry woods, Highland, 580, August 18, 1892.
(Mixed with the preceding and hence numbered the same)

LESPEDEZA INTERMEDIA S. Wats. (Hopkins, Rhodora 37: 264-266, 1935).
Dry thickets, Libertyville, 583, August 29, 1900

LeESPEDEZA StuveI Nutt. Dry woods, Highland, 582, September 12, 1896

Victa Cracca L. Roadside, Highland, 646, June 17, 1900

LATHYRUS PALUSTRIS L., var. MYRTIFOLIUS (Muhl.) Gray. Wet soil near
Libertyville, 571, June 28, 1897

LINUM VIRGINIANUM L. Field near Highland, 391, August 20, 1893

OXALIS VIOLACEA L. Moist woods near Poughkeepsie, Dutchess County, 409,
May 12, 1896

187

ERoDIUM CICUTARIUM (L.) L’Her. Weed in a cemetery, Highland, June 20,
1907

PoLyGALA SENEGA L. Woods east of Poughkeepsie, Dutchess County, 482,
June 12, 1896

EUPHORBIA PLATYPHYLLA L. Weed in cultivated field, Highland, 2370, July 21,
1892

CALLITRICHE PALUSTRIS L. Shallow water, Ohioville, 837, September 13, 1898

CALLITRICHE HETEROPHYLLA Pursh. Shallow water, Highland, 836, Septem-
ber 6, 1895

ILEx MONTANA Torr. & Gray. Thickets near Mohonk, 421, May 25, 1896

ILEX LAEVITATA (Pursh) Gray. “Pine Hole’ swamp, 2 miles south of Ohio-
vile, 419, June 3, 1896

Vitis Lasprusca L. Thickets near Highland, 443, June 6, 1893

Histscus Trionum L. Adventive, Highland, 367, October 7, 1893

HYPERICUM GENTIANOIDES (L.) B.S. P. Dry soil near Highland, 350, Septem-
ber 11, 1892

VIOLA LANCEOLATA L. Shores, Mohonk, 240, June 20, 1893

VIOLA CANADENSIS L. Woods near Claryville, 235, May 12, 1897

CUPHEA PETIOLATA (L.) Koehne. Meadow near Highland, 855, August 17,
1892

OENOTHERA GRANDIFLORA Ait. Highland, 893, July 8, 1899. Not stated whether
an escape or not. Sepals 4.5-4.75 cm. long; petals 4-4.5 cm. long; style
3.5 cm. long; stigmas spreading, about 7 mm. long

PROSERPINACA PALUSTRIS L. Wet places, Highland, 847, July 27, 1893

ZIZIA CORDATA (Walt.) DC. Dry woods, Highland, 998, May 29, 1898

HERACLEUM LANATUM L. Open moist places, Highland, 963, June 16, and
August 31, 1896

ANGELICA VILLOSA (Walt.) B. S. P. Woods near Highland, 934, August 31,
1892

CLETHRA ALNIFOLIA L. Lake Minnewaska, 1627, July 21, 1895

CHIMAPHILA MACULATA (L.) Pursh. Dry woods, Highland, 1623, July 8,
1893

PyrROLA SECUNDA L. Woods near Highland, 1663, June 18, 1893

RHODODENDRON CANADENSE (L.) B. S. P. (Rhodora canadensis L.) Minne-
waska, 1671, May 8, 1896

RHODODENDRON MAXIMUM L. Woods near Minnewaska, 1669, June 29, 1893

KAtMtIA Po.tFortaA L. “Pine Hole’ swamp, 2 miles south of Ohioville, 1638,
May 29, 1899

ANDROMEDA GLAUCOPHYLLA Link. Borders of Saxton’s Pond, Lloyd, 1616,
May 1, 1895

CHAMAEDAPHNE CALYCULATA (L.) Moench. Borders of Saxton’s Pond, Lloyd,
1621, May 1, 1895

CHIOGENES HISPIDULA (L.) Gray. Mossy woods, Claryville, 1625, May 12,
1897

VACCINIUM MACROCARPON Ait. Borders of Saxton’s Pond, Lloyd, 1685,
June 20, 1895

188

SAMOLUS PARVIFLORUS Raf. (S. floribundus H. B. K.). Wet meadow,
Libertyville, 1714, June 28, 1897

LysIMACHIA VULGARIS L. Damp soil, Ohioville, 1711, June 29, 1893

FRAXINUS NIGRA Marsh. Wet woods, Highland, 1736, May 12, 1893

GENTIANA QUINQUEFOLIA L. Woods near Libertyville, 1802, September 28,
1896

GENTIANA CLAUSA Raf. Damp thickets, Highland, 1794, September 26, 1893

BARTONIA VIRGINICA (L.) B. S. P. Mossy woods near Highland, 1784, July 21,
1900

MENYANTHES TRIFOLIATA L. Borders of Saxton’s Pond, Lloyd, 1809, May 13,
1896

ASCLEPIAS TUBEROSA L. Dry soil, Millburn, 1765, August 5, 1903

ASCLEPIAS PURPURASCENS L. Open woods, Highland, 1759, July 3, 1893

ASCLEPIAS QUADRIFOLIA Jacq. Dry woods, Highland, 1760, June 12, 1893

CONVOLVULUS SPITHAMAEUS L. Field near Butterville, 1892, May 25, 1896

Cuscuta EritrayMum Murr. On red clover, Trifolium pratense L. near
Highland, 1900, August 22, 1896

VERBENA STRICTA Vent. Field near Highland, 2073, July 23, 1902

AGASTACHE NEPETOIDES (L.) Kuntze. Woods near Highland, 2105, Septem-
ber 7, 1899

SCUTELLARIA PARVULA Michx. Slope near New Paltz, 2167, June 6, 1903

Leonurus MARRUBIASTRUM L. Waste ground, Highland, 2076, October 4,
1897

MELISSA OFFICINALIS L. Waste ground near Highland, 2113, September 7,
1892

BLEPHILA HIRSUTA (Pursh) Benth. Dry soil, Hackensack road near Pough-
keepsie, Dutchess County, 2078, July 26, 1899

MENTHA ALOPECUROIDES Hull. Waste ground near Highland, 2127, Septem-
ber 16, 1893

SoLANUM ROSTRATUM Dunal. Waste ground, Highland, 1939, September 25,
1894

PHYSALIS PHILADELPHICA Lam. (P. subglabrata Mack. & Bush). Moist field
near Highland, 1929, August 25, 1892

PHYSALIS HETEROPHYLLA Nees, var. NYCTAGINEA (Dunal.) Rydb. Meadow
near Highland, 1929, August 25, 1892

DaturA TaTuLa L. Waste ground, Highland, 1918, August 28, 1893

Mimutus aLatus Ait. Wet soil, Libertyville, 1991, July 29, 1898

LINDERNIA DUBIA (L.) Pennell (Jlysanthes dubia Barnh.). Muddy shores,
Libertyville, 1982, July 21, 1895

VERONICA ANAGALLIS-AQUATICA L. Wet soil, Highland, 2021, October 20,
1892

VERONICA LONGIFOLIA L. Established, near Lloyd, 2029, July 16, 1903

AUREOLARIA PEDICULARIA (L.) Raf. Woods near Highland, 1965 August 22,
1892

AUREOLARIA FLAVA (L.) Pennell. Woods near Highland, 1960, July 15, 1893

GERARDIA PURPUREA L. “Pine Hole” swamp, 2 miles south of Ohioville, 1966,
September 18, 1896

189

CASTILLEJA COCCINEA L. Meadow near Libertyville, 1946, May 16, 1899

UTRICULARIA CORNUTA Michx. Borders of Saxton’s Pond, Lloyd, 2041, July 8,
1895

UtTRICULARIA JUNCEA Vahl. “Pine Hole” swamp, 2 miles south of Ohioville,
2042, September 5, 1908

CONOPHOLIS AMERICANA (L.) Wallr. Woods near Highland, 2034, July 17,
1894

PLANTAGO CORDATA Lam. Hudson River near Highland, 2191, July 1, 1893

Diopta TERES Walt. Dry soil, Highland, 225, August 10, 1907

HOoUuSTONIA PURPUREA L. Meadow near Highland, 1082, June 25, 1896

LoNICERA CaApRIFOLIUM L. Naturalized near Highland, 1028, June 13, 1893

VIBURNUM PRUNIFOLIUM L. Thickets, Highland, 1050, May 26, 1893

DiIpsacus SYLVESTRIS Huds. Waste ground, Highland, 1011, August 18, 1893

SPECULARIA PERFOLIATA (L.) A. DC. Dry slopes near Highland, 1611,
June 28, 1893

CAMPANULA ULIGINOSA Rydb. Marsh near Highland, 1603, July 14, 1894

Lopet1a DortTMANNA L. Shallow water, Minnewaska, 1590, July 21, 1895

VERNONIA NOVEBORACENSIS Willd. Wet soil near Highland, 1580, August 18,
1892

EUPATORIUM SESSILIFOLIUM L. Woods near Highland, 1344, August 11, 1899

MIKANIA SCANDENS (L.) Willd. Thickets along Hudson River near’ High-
land, 1446, August 28, 1892

SoLipaco PATULA Muhl. Marsh near Highland, 1531, November 7, 1892

SoLipAco oporA Ait. Minnewaska, 1529, July 28, 1896

SoLmpAGO NEGLECTA Torr. & Gray. “Pine Hole’ swamp, 2 miles south of
Ohioville, 1525, September 18, 1896

SOLIDAGO ULMIFOLIA Muhl. Thickets near Highland, 1551, September 26,
1898

AstTER Hervey Gray. Woods near Highland, 1171, August 20, 1899

AsTER TRADESCANTI L. Wet soil near Highland, 7158, October 16, 1892

ASTER PANICULATUS Lam., var. ACUTIDENS Burgess. Dry soil near Highland,
1192, October 18, 1892

ASTER PUNICEUS L., var. FIRMUS (Nees) T. & G. Damp soil near Highland,
1185, October 25, 1894

ASTER LINEARIFOLIUS L. Dashville Falls, 1175, October 11, 1901

ASTER INFIRMUS Michx. Open woods, Highland, 1172, July 26, 1893

RUDBECKIA SPECIOSA Wenderoth. New Paltz, 1476, September 6, 1899.
Probably an escape.

CorEOPSIS TINCTORIA Nutt. Highland, 1288, August 22, 1893. Not stated
whether cultivated, naturalized or an escape.

BIDENS coMosA (Gray) Wiegand. Wet soil, Highland, 1232, August 31, 1892

BIDENS BIPINNATA L. Moist ground, Highland, 1231, August 28, 1892

BIDENS CERNUA L. Marsh near Milton, 1233, September 15, 1896

This is the tall, erect form with rays exceeding the bracts and with

elongated narrow leaves, prominently and distantly toothed. Being essen-
tially glabrous it is often confused with Bidens laevis. It is the common

form of this species in the marshes of the Hudson River and adjacent
territory, and rarely elsewhere across the state.

190

BIDENS TRICHOSPERMA (Michx.) Britt. Saxton’s Pond, Lloyd, 1289, Septem-
ber 5, 1895. Possibly native, although the same plant has appeared at
several localities in eastern New York within recent years, and evidently
introduced

CHRYSANTHEMUM PARTHENIUM (L.) Bernh. Adventive, Highland, 1258,
September 10, 1892

SENECIO oBpovATUS Muhl. Woods near Highland, 1482, May 25, 1896

Cirsium piscotor Muhl. Open woods near Highland, 1294, September 15,
1892

CirsIuUM MuTICUM Michx. Swamp near Highland, 1298, October 11, 1893

CirsIuM PUMILUM (Nutt.) Spreng. Dry fields, Highland, 1300, August 26,
1892

CENTAUREA JACEA L. Waste ground, Highland, 1253, August 7, 1903

LAPSANA COMMUNIS L. Waste ground near Highland, 1425, June 21, 1897

KRIGIA BIFLORA (Walt.) Blake (K. amplexicaulis Nutt.). Weed in cultivated
soil, Highland, August 8, 1923

KRIGIA VIRGINICA (L.) Willd. Dry soil, Highland, 1412, June 7, 1894

VIREA AUTUMNALIS (L.) S. F. Gray (Leontodon autumnalis L.). Roadside
near Highland, July 20, 1907

PIcRIS HIERACIOIDES L. Waste ground near Highland, July 10, 1907

Picris ECHIOIDES L. Roadside near Highland, July 10, 1907

LACTUCA CANADENSIS L., var. OBOVATA Wiegand. Woods near Highland,
1419, July 15, 1903

HIERACIUM PRATENSE Tausch. Fields near Highland, July 10, 1907

HieRAcIuM Gronovit L. Open woods near Highlands, 1396, August 22, 1895

HIERACIUM CANADENSE Michx. Dry woods, Libertyville, 1393, August 30,
1898

New York STaTE MusEuM
ALBANY, N. Y.

BOOK REVIEWS
Flora of Fukien

Flora of Fukien and Floristic Notes on Southeastern China. First Fas-
cicle. By Franklin P. Metcalf, ix. + 82 double pages. Lingnan University,
American Office, 150 Fifth Ave., New York City. $1.50.

This is the first part of a monumental work on the flora of
southeastern China. Dr. Metcalf has already given twenty years
to the task. He served as Professor of Botany in the Fukien Chris-
tian University from 1923 to 1929, and in the Lingnan University
in Canton (formerly Canton Christian College) from 1931 to 1938.
He and his students have collected extensively in China, and a
Rockefeller fellowship made it possible for him to see practically all
of the Fukien plants in the herbaria of the world. Since leaving

191

China he has been giving his whole time to this flora working with
Dr. E. D. Merrill at the Arnold Arboretum.

According to Dr. Metcalf there is no book or group of books
by which the plants of southeastern China can be identified and
specimens have to be sent to specialists in Europe or America. This
work will be a landmark in Chinese botany. Bentham’s Flora Hong-
kongenis was published in 1861 and is out of date and out of print,
besides covering only a small area.

This fascicle covers fourteen families from Cycadaceae to
Fagaceae. There are keys to families, genera and species. There is
a description of each Fukien species and additional notes on those
found in adjacent provinces. No new species are described but at-
tention is called to many novelties which are to be described later.

It is hoped that future parts can be published in China but the
war made it necessary to publish this fascicle here. This work is
another reminder of the many contributions of missionaries to Chi-
nese botany. Hundreds of species were first sent to western botan-
ists by friars, abbés and clergymen. In recent years good collections
were being built up in the Chinese colleges until the Japanese inter-
fered.

My only criticism is that in the interest of economy the page
margins are very narrow and there is little room for additional
notes. I should think that one using it very much would have to
have it interleaved. R. R. Stewart

Botanizing in Cuba

Itinéraires botaniques dans l’ile de Cuba. (Premiére série). By Frére
Marie-Victorin, F.E.C., D.Sc., Directeur de l'Institut botanique de 1’Univer-
sité de Montréal, and Frére Léon, Directeur du Laboratoire de botanique
du Colegio de la Salle, Havana. Contributions de l’Institut botanique de
l'Université de Montréal, No. 41. Montréal, 1942.

The writer of this review was companioned, on what was for
us a botanizing trip from Capetown through Egypt to Jerusalem
and farther, by the senior author of the above publication. It was
immediately apparent that Frére Marie-Victorin was an insatiable
diary keeper, an amiable weakness I thought! Until a copy of this
diary, beautifully bound, was put into my hands some time after
our return to Montreal, I confess I did not realize that weakness
had become strength. Here before me was a volume almost fit for
publication, displaying a general picture of the vegetation and its

192

habitants, and a good many other matters not usually regarded as
botanical, but enriching the picture from the human point of view.
Very much such a work is that now lying before us. We well know
that Marie-Victorin is enamoured of Cuba and has been busy for
some time, in cooperation with Frere Léon, in studying its vegeta-
tion. But the results of their work embrace more than collections
and descriptions of long lost or new species. One of these results
is this volume of “Itineraries” by the perusal of which botanists
interested in tropical vegetation (and what botanist is not?) will
gain a vivid impression of what may be seen in the island of Cuba.
This is the more so because of the plenitude of illustration. As the
reviewer knows, the camera is almost a part of Marie-Victorin, and
we see in this publication embellished with about 280 photographs,
with a number of line drawings and a large map, an account which
appeals directly to the eye. Thus one gets a full and detailed impres-
sion of how the country actually looks, and one feels as if he had
seen Cuba for himself. The liberal use of native names and frequent
descriptions of the uses made of the vegetable products enhances
this impression. More than this, there are many allusions to human
relations, some of which appeal directly to the heart.

Francis E. Lioyp

Diary and Travels of the Bartrams

Diary of a Journey through the Carolinas, Georgia, and Florida from
July 1, 1765 to April 10, 1766. By John Bartram. Annotated by Francis
Harper. Transactions of the American Philosophical Society 33(1) : iv + 120.
portrait, 8 maps, 37 fig. December, 1942. Paper cover $2.00.

Travels in Georgia and Florida, 1773-74; A Report to Dr. John Fother-
gill. By William Bartram. Annotated by Francis Harper. Trans. Am. Phil.
Soc. 33(2): about 115 pp. portrait, 5 maps, 47 fig. Spring 1943. Paper
cover $2.00. Parts 1 and 2 bound together in cloth $5.00.

John Bartram was a Quaker botanist to the King of England.
He was the first botanical investigator of the upper reaches of the _
St. John’s River in Florida, and of the greater part of Georgia.
As a friend and guest of the élite in Charleston, and of several
colonial governors, he observed and portrayed pre-Revolutionary
life in the southern cities of Charleston, Savannah, and St. Augus-
tine, as well as life on the plantations and in the wilderness. He
described the architecture of the first Spanish period in St. Augus-
tine, and told of the easternmost known calumet ceremony, at

193

the Treaty of Picolata. On this trip he discovered Franklinia,
Pinckneya, Nyssa ogeche, Canna flaccida, and other noteworthy
plants. He was accompanied and assisted by his talented son,
“Billy,” who was destined to become the author of the immortal
Travels (1791).

The record of these achievements appears in the simple, un-
varnished diary of 1765-66, preserved at the Historical Society
of Pennsylvania and hitherto largely unpublished. The full editorial |
comments and annotations provide a historical background, identify
Bartram’s plants and animals, and show his routes in detail by
means of both colonial and modern maps. Photographs and draw-
ings bring into vivid focus, after a span of nearly two centuries,
many of the points of particular interest that were visited by John
Bartram.

Much new light on William Bartram’s celebrated Travels
(1791) will be forthcoming with the publication, in part 2, of
his lengthy manuscript report to his London patron, Dr. John
Fothergill. This important document, which has long remained in
obscurity in the British Museum, will be a distinct boon to all
students of Bartram and of early American natural history. While
it covers the same ground as the first part of the book of 1791, it
is not a duplicate of that work, but contains much additional
information on Bartram’s itinerary, his chronology, his scientific
and literary qualifications, and the identification of his plants and
animals. The work is thoroughly annotated and indexed. The
illustrations include the most significant collection of Bartram
drawings ever brought together in a single publication.

HUNTER COLLEGE Haroitp H. CLum
New York, N. Y.

Carnivorous Plants

The Carnivorous Plants. By Francis Ernest Lloyd. xv + 352 pages, 38
plates. Waltham, Mass., The Chronica Botanica Co., New York City, E. G.
Stechert and Co. 1942. $6.00.

“For the present moment, I care more about Drosera than the
origin of all the species in the world,’ wrote Charles Darwin, in
1860, to his friend Sir Charles Lyell. It is fortunate that he found
time to consider both, for his investigations on Dionea, Drosera,
and physiologically related plants resulted in the publication of the

194

first book on this group, which has remained the only one available
in English for the past sixty-eight years.

It is in a similar spirit that Dr. Lloyd has now summarized the
researches in this field. Karl von Goebel reviewed the work in 1891
in a section of his book, “Pflanzenbiologische Schilderungen.”
Since then, various contributions have been made, including the
skillful researches of Lloyd on Utricularia and the respective studies
of Vines and Hepburn on the digestive action of the pitcher fluids
of Nepenthes and Sarracema. The glandular secretions of Drosera
were investigated in a similar manner by Okahara. The existence
of carnivorous fungi has recently been established, and new inter-
esting interpretations of the mechanisms of closure of the leaves of
Dionea and Aldrovanda have been made by Brown in the United
States and Ashida in Japan.

There has long been a need for a comprehensive, modern treat-
ment of the carnivorous plants (Dr. Lloyd prefers this term to in-
sectivorous). The present volume seems entirely adequate. It
is a precise and scholarly work. The author has carried on in-
tensive investigations in this field since 1929, when he made his
first observations on the trap of Utricularia gibba. The book con-
tains a great stock of his own experiments and verifications of the
results of others. For example, in the case of Roridula, Dr. Lloyd
is now definitely able to exclude this genus from the carnivorous
plants. Many of the plants, especially the Utriculariae, were studied
in their native habitats. Two trips to Africa and one to Australia
were made for this purpose.

The text is divided into fourteen chapters, each for the most
part corresponding to a separate genus. The distribution of the
plants is unusual. They either fall into groups which are widely dis-
tributed, like Drosera or Utricularia, or else they exist as monotypic
or very local genera, as Cephalotus, Genlisa, Dionea, and others.
The chapters are arranged according to the increasing order of com- |
plexity of the traps. Thus we find, in this rather ingenious system
of classification, that the passive, pitfall traps, as represented by
the pitchered leaves of Heliamphora and Sarracenia, are placed first.
Passing upward through the lobster pots, snares, fly-paper and
active, bear-trap devices, we come to the mouse trap, or most com-
plex type which includes such forms as Biovularia, Utricularia,
and Polypompholyx. One chapter is devoted to the fungi that

195

prey upon small water animals, notably eelworms and rotifers.
Modifications encountered in this unique group are hyphal loops
that swell and clamp onto the body of a worm unluckly enough to
enter, and sticky plugs that literally gag their victims to death. This
appears to be the first complete discussion of the findings of
Drechsler and others on these fungi. Although Dr. Lloyd, in re-
viewing the genus, mentions the use of the name “Chrysamphora” in
place of the much more familiar “Darlingtoma,’ which it antedates,
he continues to use the latter name. The literature cited on Utricu-
laria records twelve papers by the author embodying new concepts
on. the operation of the traps, and an additional two describing four
new species discovered in Australia. An appendix is added as a sort
of patent registry to describe an epoch making mouse catching de-
vice constructed on the principles of the Utricularia trap. Its efficacy
is assured, since 1f it did not catch the mouse it would undoubtedly
leave him with a severe nervous breakdown.

With very few exceptions, the drawings are original. They are
ample and very well done. The plates are included at the back. Due
to a regrettable economy of space, many of the photographs have
been cut down and placed in a sort of mosaic on the page, with a
resultant loss of clarity. In every other detail, however, the book
shows great care in its preparation. The paper, print, and binding
are good. The frontispiece is an old drawing of a species of Nepen-
thes from an early herbal.

Above all, it is well to remember that ““The Carnivorous Plants”
represents the achievement of more than twelve years of painstaking
work. Dr. Lloyd has not left the smallest pebble unturned in his
effort to follow up every source. As witness to his consuming in-
terest in the field, there are very few chapters that do not contain
some of his own pertinent, and usually outstanding, contributions.

CoLuMBIA UNIVERSITY Oscar J ANIGER

RIDBILID) INNUES OUP Wale, CILAWs}

TRIP OF OcTOBER 4, 1942, To RicHMonp, S. I.

Nine members of the Club took this trip. The main objective
was the salt marsh, but many interesting plants were seen along
the road on the way to the salt marsh. Over a hundred species were
pointed out, and about as many more were passed by without men-
tion because they were so familiar. Several members of the group
besides the leader were alert in spotting plants, and helpful in
identifying them and in making a list, as well as in finding the
way out of the woods after leaving the salt marsh.

Some of the easily recognized grasses promised by the Field
Chairman were the tall and spreading switch grass (Panicum.
virgatum), the always interesting hispid panicum (P. clandes-
tinum), the very delicate old-witch grass (P. capillare), tall red
top (Tridens flavus) with its purplish glumes that rub off black,
the graceful and silky Indian grass (Sorghastrum nutans), the
large coarse gama grass (Tripsacum dactyloides) with its polished
jointed spikes, wild rye (Elymus virginicus), which, like a cat,
resists being petted the wrong way, broom beard grass (Andro-
pogon scoparius) with its spreading feathery hairs, Virginia beard
grass (A. virginicus) and its bushy-headed form A. glomeratus,
the delicate but savage rice cut-grass (Leersia oryzoides), and the
bristly-sheathed salt-marsh cockspur grass (Echinochloa Walteri).

In or near a brook or road-side ditch were found, not in bloom,
water-weed (Elodea canadensis), water starwort (Callitriche palus-
tris), sweet flag (Acorus Calamus) and the hairy variety of swamp
milkweed (Asclepias incarnata var. pulchra), the last in fruit.

Some of the less common trees and shrubs along the road were
two somewhat southerly species, clammy locust (Robima viscosa)
and false indigo (Amorpha fruticosa), the latter in fruit ; the middle
western Osage orange (Maclura pomifera) in fruit; also hack-
berry (Celtis occidentalis) in fruit, box elder or ash-leaved maple —
(Acer Negundo), and a particularly large specimen of tulip tree
(Liriodendron Tulipifera).

A high point on the hill, to which the road led, afforded a good
general view of the salt marsh and its creek, with the character-
istic winding or crooked form for which genuine creeks like this
are named.

196

197

Eleven species of aster (including A. paniculatus and two of
its varieties) were seen besides New York aster (A. novi-belgi)
and the two salt-marsh asters A. subulatus and A. tenuifolius.
Six species of goldenrod were found besides seaside or salt-marsh
goldenrod (Solidago sempervirens ).

In the moist ground near the salt marsh were found swamp
thistle (Cirsium muticum), rough thoroughwort (Eupatorium ver-
benaefolium), soapwort gentian (Gentiana Saponaria), tall sun-
flower (Helianthus giganteus), ladies’ tresses (Spiranthes cernua),
Culver’s-root (Veronica virginica) in fruit, and the tiny water
pimpernel (Samolus floribundus).

Of the real salt-marsh plants the most interesting, besides the
few already mentioned, were the two shrubby composites, ground-
sel tree (Baccharis halimifolia) with beautiful plumy white heads
on the pistillate plants, and the less showy marsh elder (Iva oraria),
the red patches of the fleshy glasswort (Salicorma europaea), the
extensive wiry carpet of the so-called black grass (Juncus Gerardi),
the dioecious alkali grass (Distichlis spicata), the pink-flowered
salt-marsh fleabane (Pluchea camphorata) with its characteristic
aroma, the beautiful and delicate marsh pink (Sabatia stellaris),
the tall weak unattractive water hemp (Acnida cannabina), orach
(Atriplex patula var. hastata) turning red in places, beaked spike
rush (Eleocharis rostellata) looping its way along, salt-marsh bul-
rush (Scirpus robustus), Olney’s bulrush (S. Olneyi), the low
cord grass (Spartina patens), and the tall salt-marsh grass (S.
glabra var. alterniflora—nomenclature of Gray’s Manual used here
and throughout).

Two characteristic plants known to grow in this salt marsh,
but not seen by the group on this trip were a third species of
Spartina, salt reed grass (S. cynosuroides), and the lovely sea
lavender (Limonium carolinianum).

Hester M. Rusk

Trip oF NovemMBer 15, 1942, to Laxewoonp, N. J.

The walk included an old pine barrens bog, dry barrens, and
the lake shore. The leader pointed out plants typical of the habitats
and others of interest. The most important discovery was made
by Mr. A. T. Beals,.a moss which was finally identified by Dr.
Grout as Entodon seductrix var. minor (Aust.) Grout. Mr. Beals

198

writes in part, “This variety is a find (for New Jersey). It is not
included in any list I have seen of New Jersey mosses, although
it may have been collected previously in that state. The plant is
more common further south and was named and described from
a specimen found in Georgia.”

In the late afternoon Mr. V. L. Frazee arranged for us to visit
a Mr. Lecompte who is related to Dr. Knieskern. We saw some of
Knieskern’s collections.

Attendance: 24. Leader: Mr. James Murphy.

PROCEEDINGS OF THE CLUB
MINUTES OF THE MEETING OF NOVEMBER 2, 1942

The meeting was called to order at 8:25 p.m. by the President,
Dr. C. Stuart Gager, at the Museum of Natural History. Thirty-
two members and friends were present.

The minutes of the preceding meeting were accepted as read.

The election of Mr. Mario G. Ferri, Departamento de Botanica,
Faculdade de Fiolsofia, Ciencias e Letras, Caixa Postal 2926,
Sao Paulo, Brasil, to annual membership was unanimously
approved.

The suggestions proposed in the report of the Per Capita Cost
Committee were read by Dr. Matzke in the absence of the chairman
of the committee.

The scientific program of the evening was presented by Dr.
Henry K. Svenson who spoke on the “Vegetation of Western
South America.” The talk was illustrated with Kodachrome slides
which depicted the vegetation, peoples and points of interest in
that region.

The meeting was adjourned at 9:35 p.m.

Respectfully submitted,

Honor M. HoL_INnGHURST
RECORDING SECRETARY

MINUTES OF THE MEETING OF NOVEMBER 18, 1942

The meeting was called to order at 3:30 p.m. by the second
Vice-president, Dr. Clyde Chandler in the Members Room of the

199

New York Botanical Garden Museum. Twenty-nine members and
friends were present.

The minutes of the preceding meeting were accepted as read.

The first part of the scientific program was presented by Mrs.
Annette Hervey who spoke on “The Use of Phycomyces Blakes-
leeanus in the Assay of Thiamin in Agar.” The talk was illustrated
with slides.

The second portion of the program was presented by Mr.
John D. Dodd who gave an illustrated talk on “Three Dimensional
Cell Shape in the Carpel Vesicles of Citrus Grandis.” The speak-
er’s abstract follows:

Internal cells from the carpel vesicles (juice sacs) of grapefruit were
examined in the living condition. Cell walls were stained lightly with neutral
red. Records were kept by making a careful drawing of each cell. In order
to insure completely impartial selection, the data were not tabulated and
summarized until 100 cells had been drawn. Results showed an average of
13.85 faces per cell. The range in number of faces was from 9 to 18.
The largest number of any one type was 22 cells each with 14 faces. Of
the rest 39 cells had more than 14 faces and 39 had less. The number of
edges per face varied from 3 to 8; 0.8% were triangular, 25.9% were

quadrilateral; 41.6% were pentagonal; 23.6% were hexagonal; 7.0% were
heptagonal and 1.1% were octagonal.

The meeting was adjourned at 4:30 p.m. and was followed by
tea which was served by friends at the Garden.
Respectfully submitted,

Honor M. HoLttincHurRST
RECORDING SECRETARY

MINUTES OF THE MEETING OF DECEMBER 1, 1942

The meeting was called to order at 8:25 p.m. by the President,
Dr. C. Stuart Gager, in the Museum of Natural History. Thirty-
five members and friends were present.

The minutes of the preceding meeting were accepted as read.

The scientific program of the evening was presented by Mr.
Otto Degener who gave an illustrated talk on “Botanizing in Fiji.”

The speaker’s abstract follows:
; While a member of the Pacific cruise of the palatial junk-yacht, “Cheng-
Ho,” sponsored by Mrs. Ann Archbold, collections were made, under the

auspices of the Arnold Arboretum and the New York Botanical Garden,
amounting to about 2,100 numbers or a total exceeding 15,000 specimens.

200

These are being studied by Dr. A. C. Smith and various specialists. Thus
far 64 novelties have been described, one proving to belong to an entirely
new family related to the Magnoliaceae, Himantandraceae and Winteraceae.
Mr. Degener, with the aid of his “adopted Figi son’ Aloisio (Aloysius)
Tabualewa, won the confidence of the Fijians who ordinarily do not look too
kindly on the aggressive papalangi or white man, and lived with them in
their elaborately constructed “grass” houses. This enabled him to collect
data on their customs and how they used certain plants in their native medi-
cine and arts. Their use, for example, of the latex of various species of
Alstonia, as chewing gum, may help us solve the problem of soothing the
nerves of countless ruminating stenographers, should our national supply of
American chicle give out. Thirteen-year-old Leroy Peiler, a native Hawaiian
refugee and Mr. Degener’s ward, later served yangona, a beverage made from
Piper methysticum, in proper Fiji style.

The meeting was adjourned at 9:30 p.m.

Respectfully submitted,

Honor M. HoLLtinGHuURST
RECORDING SECRETARY

MINUTES OF THE MEETING OF DECEMBER 16, 1942

The meeting of December 16 was held in Larkin Hall of Ford-
ham University. Thirty-seven members and friends were present.
Preceding the regular meeting, the members of the Torrey Botanical
Club were invited to inspect the biological laboratories and to
observe microscopic demonstrations. Refreshments were then
served.

The meeting was called to order at 4:50 p.m. by the President,
Dr. C. Stuart Gager, who introduced the first speaker, Father
Berger of Fordham University. The topic presented by Father
Berger and Miss Eleanor Witkus was “The Prophases of Poly-
somatic Mitosis and their Relation to Meiosis.” The speakers’
abstract follows:

The essentials of Darlington’s precocity theory of meiosis, the singleness:
_ of leptotene chromonemata, the attraction in pairs only and the repulsion
between pairs of pairs, and metaphase pairing due to chiasmata, were pre-
sented and refuted in the light of evidence brought forward by our spinach
material and the work of other investigators.

In the periblem of the root tips of Spinacia oleracea, in addition to diploid
cells with twelve chromosomes, tetraploid and octoploid cells are regularly

201

found. This condition of polyploidy arises by double chromosome reproduc-
tion during the resting stage. In the prophase and metaphase of certain of
these polysomatic cells the chromosomes are in closely associated pairs.

In Spinacia oleracea, therefore, more than two chomonemata may be
present in closely paired association. In such multiple associations there is
no evidence of any repulsion between pairs. Paired associations are main-
tained from earliest prophase to metaphase without being held together by
chiasmata.

After the discussion of these talks, Dr. Gager expressed the
thanks of the Torrey Botanical Club to Father Berger and his staff
for their kind hospitality. The meeting was adjourned at 5:45 p.m.

Respectfully submitted,

Honor M. HoLitincHurst
RECORDING SECRETARY

DATES OF PUBLICATION OF TORREYA, VOLUME 42

Number 1. January-February February 27, 1942
2. March-April ; April 10, 1942
3. May-June June 5, 1942
4. July-August November 12, 1942
5. September-October January 29, 1943
6. November-December April 24, 1943

ERRATA

Page 56, line 6 from bottom: for nutalii, read Nuttallii.
Page 66, line 7 from bottom: for Dr. Wm. J. Crocker, read Dr.
Wm. Crocker.
Page 73, line 4 from bottom: for datas, read data.
Page 97, first new member listed: for Miss Marion Johnson, read
Mr. Marion Johnson.
Page 101, fourth new member listed: should be Professor Hemp-
stead Castle, Yale University.
Page 126, bottom line: for Erythina, read Erythrina.
Page 129, for Rynchosphora, read Rynchospora.
for Scirpa, read Scirpus.
for Schleria, read Scleria.
Page 143, line 5: for Botrichium, read Botrychium.

INDEX TO TORREYA—VOLUME 42

New names are in bold face type

Abies Webbiana, 100

Achras calcicola, 69, 71; chicle, 69-72,
74; gapota, 38, 42, 69-81, 105, 110,
111

Acronychia, 126

Adenanthera pavonina, 126

Adlumia fungosa, 185

Agastache nepetoides, 188

Aglaia samoensis, 126

Agrostemma Githago, 185

AJELLo, Lipero. A new Chytrid
genus Polychitrium, lecture, 145

Alaska, The genus Cicuta in, 176

Aleurites moluccana, 127

Allegany County, N. Y., 174

Allegheny Plateau, 174

Arren 1G. E65

Alphitonia zizyphoides, 126

Alstonia eximia, 77, 81; grandiflora,
77, 81; Scholaris, 77, 81

Alyxia stellata, 126

Amanita muscaria, 84, 85; Peckiana,
83

Amanitopsis vaginata var. plumbea,
82; volvata, 82

Anacharis occidentalis, 183

Anchistea virginica, 182

Anperson, J. P. Flora of Alaska—
I. The genus Cicuta, 176

ANDERSON, Louts, 60

Andromeda glaucophylla, 187

Angelica villosa, 187

Aplectrum hyemale, 184

Arabis canadensis, 185

Arenaria groenlandica, 185; lateri-
flora, 185

Arethusa bulbosa, 184

Arisaema Dracontium, 184; pusillum,
183

Aristolochia Serpentaria, 184

Arizona, flora of, 9, 138

Asarum canadense var. acuminatum
f. Prattii, 180; caudatum f. chlo-
roleucum, 180

Asclepias purpurascens, 188; quadri-
folia, 188; tuberosa, 188

Asplenum auritum, 34; exiguum,
139; montanum, 182; pinnatifidum,
144: trichomanes, 95

Aster Herveyi, 189; infirmus, 189;
linearifolius, 189; paniculatus, 189;
puniceus, 189; Tradescanti, 189

Atherium alpestre var. americanum,
145

Aureolaria flava,
188

Autumn coloration, lecture, 27

188; pedicularia,

Bald cypress, 57

Batpwin, H. I., 23, 24

BarNnuHArpt, J. H., elected delegate,
30

Barringtonia asiatica, 127

Barton, L. V. Some special prob-
lems of seed dormancy, lecture, 63

Bartonia virginica, 188

Befaria, 168

BENEpDIcT, R. C., 23

BenHAM, RuHopA W. The microbe’s
challenge, review, 88

Benson, Lyman. Notes on the flora
of Arizona, 9

Benthamia, 11, 12; Spach, 12

Benthanudia, 11

Bercer, C. A. and ELeanor WIrt-
Kus. The prophases of polysomatic
mitosis and their relation to meio-
sis, lecture, 200

Berteroa incana, 185

Betula Bhojpattra, 100; nigra, 184

Bidens bipinnata, 189; cernua, 189;
comosa, 189; trichosperma, 190

Bikkia grandiflora, 126

Billbergia, 96

BLAKESLEE, A. F., 67

Blechnum, 36; spicant, 145

Blephila hirsuta, 188

Botp, Harotp C. An introduction to
the study of Algae, review, 140

202

203

Boletus castaneus, 84; chrysenteron,
84; granulatus, 84
BonistEeEL, W. J. Hunger signs in
crops, review, 21; electe’ to Coun-
cil, 63; resigned as editor of Tor-
REYA, 103
Book Reviews:
Chapman. An introduction to the
story of Algae, 140
Deam. Flora of Indiana, 53
Eberson. The microbe’s challenge,
88
Fischer & Harshbarger. The flower
family album, 51
Foster. Practical plant anatomy, 88
Fuller. The plant world, 18
Harper. Diary and travels of the
Bartrams, 192
Hayes & Immer., Methods of plant
breeding, 137
Kearney & Peebles. Flowering
plants and ferns of Arizona, 138
Kelsey & Dayton. Standardized
plant names, 132
King. Bible plants for American
gardens, 114
Large. The advance of the Fungi,
19
Larkey & Pyles. An herbal (1525),
91
Lloyd. The carnivorous plants, 193
Marcy & Shepard. Butterflies, 52
Marie-Victorin & Léon. Itinéraires
botaniques dans 1’ile de Cuba, 191
Metcalf. Flora of Fukien, 190
Needham. About ourselves, 86;
Introducing insects, 50
O’Hanlon. Fundamentals of plant
science, 90
Rodgers. John Torrey, 135
Sampson. Work book in general
botany, 115
Symposium, Hunger signs in crops,
21
Botrychium cicutarium, 37; lanceo-
latum var. angustisegmentum, 143,
182; matricariaefolium, 143; multi-

fidum var. silaifolium, 182; under-
qwoodianum, 37

Branch Brook Park, Newark, 143

Bromus hordaceus, 183

Brooklyn Botanic Garden, 58, 59

Brosimum alicastrum, 75; utile, 75,
78

Brown, Otway, 57

BuLLeTIN of the Torrey Botanical
Club, dedication of volume 69, 61

Bumelia Guatemalensis, 74;
folia, 75

Bunchberries, 11, 12

Bushkill Falls, Pa., 141

Carn, S. A., 67

Calamagrostis cinnoides, 183

California poppy, 10

Callitriche heterophylla, 187; palus-
tris, 187

Calocarpum mammosum,
viride, 72, 74

Calophyllum inophyllum, 127

Cameraria belizensis, 76

Camp, W. H. Work book in general
botany, review, 115; The genetic
structure of populations and the
delimitation of species, lecture, 147;
and C. L. Gutty. Polypetalous
forms of Vaccinium, 168

Campanula uliginosa, 189

Camptosorus rhizophyllus, 95

Canarium Harveyt, 126

Capparis sandwichiana, 126

Carex, 119, 129; aestivalis, 174, 175;

lauri-

(2a

Baileyi, 175; cephalantha, 183;
cryptolepis, 183; Davisti, 183;
exilis, 183; Grayu, 183; grisea,

183; lasiocarpa, 183; lurida, 129,
175; var. gracilis, 174, 175; planta-
ginea, 183; radiata, 174; squarrosa,
183; torta, 183
Carya alba, 184
Cassia marilandica,
186
Castilla elastica, 75, fallax, 75
Castilleja coccinea, 189

nictitans,

186;

204

Cavendishia, 34

Cell shape in the carpel vesicles of
Citrus Grandis, lecture, 199

Celtis paniculata, 126

Centurea Jacea, 190

Cephaelis, 36

Cerastium nutans, 185; oblongtfo-
lium, 185

Ceratostomella, vitamin reactions,
30-32

Cetarach dalhousiae, 139

Chamaedaphne calyculata, 187

Chamaepericylmenum, 11, 12

Cheilanthes lanosa, 144

Chicle, collecting in the American
ELODICS Heat elem son ante Oo
Part 3, 105

Chile tarweed in Quebec, 68

Chiloscyphus rivularis, 141

Chimaphila maculata, 187

China, flora of, 190

Chiogenes hispidula, 187

Chlorella vulgaris, chemical inhibi-
tion of photosynthesis in, lecture,
28

Chromosome relocation and degen-
eration in relation to growth and
hybrid vigor, lecture, 102

Chrysanthemum Parthenium, 190

Curyster, M. A. A botanist’s sum-
mer in Costa Rica, 33

Cicuta douglasii, 176-178; maculata,
176-178; mackenzieana, 176-178

Cimicifuga racemosa, 185

Cirsium discolor, 190; muticum, 190;
pumilum, 190

Cladomia pyxidata var. neglecta f.
centralis, 49; squamosa f. carneo-
pallida, 49; turgida, 49, 61

Cladoniae in New Jersey, 49

CLAUSEN, Rosert T. Carex aestivalis
and C. lurida var. gracilis on the
Allegheny Plateau, 174

Claytonia caroliniana, 185

Clematis verticillata, 185

Clethra alnifolia, 187

Clitocybe infundibuliformis, 85; pino-
phila, 85; vilescens, 85

Cium, H. H. Assumed editorship of
TorreyaA, 120; Diary and travels
of the Bartrams, review, 192

Collybia confluens, 85

Colubrina asiatica, 126

Commelina communis, 184

Conophohs americana, 189

Convallaria majalis, seed dormancy,
lecture, 64

Convolvulus spithamaeus, 188

Coptis trifolia, 142, 185

Corallorrhiza odontorhiza, 184

Cordia subcordata, 127

Coreopsis tinctoria, 189

Cornel, 11

Cornelian cherries, 11

Cornella, 13

Cornus, 11-14, 130, 131; The names
of, 11; alba, 13; canadensis, 12, 58,
143; disciflora, 13; florida, 11, 13,
14; Kousa, 13; mas, 11-13; mas-
cula, 12; Nuttallii, 13; sanguinea,
13; stolonifera, 13; suecica, 12;
V olkensiu, 13

Coronilla varia, 186

Corylus cornuta, 184

Costa Rica, A botanist’s summer in,
33

Costus, 33

Couma Guatemalensis, 76;
78; utilis, 77, 78

Crocker, W., 66

CroizaT, Leon. Phyllanthus nummu-
lariaefolius in the U. S., 14

Crotalaria anagyroides, 123; sagit-
talis, 186

Crysophyllum oliviforme, 72, 76

Cuba, Botanizing in, 191

Cuphea petiolata, 187

Cuscuta Epithymum, 188

Cyanococcus, 169

Cynoxylon, 11, 130, 131

Cyperus, 129

Cypripedium reginae, 184

Cystomium falcatum, 145

sapida,

205

Dalibarda repens, 186

DANSEREAU, PIERRE, 24

Datura Tatula, 188

DerGENER, Otto. Botanizing in Fiji,
lecture, 199

Delonix regia, 127

Delphinium Ajacis, 185

Desmodium bracteosum, 186; canes-
cens, 186; laevigatum, 186; mari-
landicum, 186; rigidum, 186

Dianthus deltoides, 185

Dicentra canadensis, 185

Dicranopteris Bancroftu, 35; costari-
censis, 35; retroflexa, 35

Diodia teres, 189

Diospyros, 126

Dipholis salicifolia, 74; Stevensoni,
72, 74

Dipsacus sylvestris, 189

Dix, W. L., 60; Rare Cladoniae in
N. J., 49; Field trip, June 12-13,
1942, 142

Dopp, J. D. Some reactions to graft-
ing in Viola, lecture, 28; Three
dimensional cell shape in the carpel
vesicles of Citrus Grands, lecture,
199

Dopcr, B. O. The advance of the
Fungi, review, 19; Hybrid vigor or
heterocaryotic vigor in the Fungi,
lecture, 149

Dodonaea viscosa, 126

Dogwood, 11, 13, 14

Dott, W. H. Field trip, June 20,
1942, 144

Dothistroma, 28

Doucias, Beaman. Botanizing in an
art museum, lecture, 142

Drimys Wintert, 36

Drosera intermedia,
folia, 185

Dryopteris, 36; celsa, 144; chinensis,
144; Goldiana, 95; Iludoviciana,
144; marginalis, 95

Dues of members in the armed forces,

149

185;

rotundi-

Dwyer, J. D. Interesting plants of
Litchfield Co., Conn., lecture, 26
Dyera borneensis, 77, 81; Costu-
lata, 77, 8 laxsfolia, 77, 81;

Lowii, 77, 81
Dysoxylum Richu, 126

Eastern New England Tour, 23

Elaeocarpus samoensis, 126

Eleocharis, 129; capitata, 183

Eleusine indica, 183

Ellatostachys falcata, 126

Elting, Clarence J., and his herba-
rium, 181

Elymus Wiegandu, 183

Englewood Cliffs, 126

Enterosora spongiosa, 34

Entodon seductrix var. minor, 197

Eriocaulon septangulare, 184

Eriophorum callithrix, 143; viridi-
carinatum, 183

Erodium cicutarium, 187

Erythrina variegata var. orientalis,
126, 127

Eschscholtzia aliena, 11; arizona,
11; californica, 10, 11; Jonesu, 11;
mexicana, 11; paupercula, 11

Eugenia, 126

Eukrania, 12, 14, 130, 131

Eupatorium angulare, $5) 8
folium, 189

Euphorbia dictyosperma, 16; platy-
phylla, 187; spathulata, 15

sessili-

Fagraea Berteriana, 126
FARWELL, OLiver A. Cornus, a reply,
130
Fassett, Norman C. Some new
_ forms from the middle west, 179
Fern garden of W. H. Dole, 144
Ficus, 126; glabrata, 76; lapathi-
folia, 76
Field Trips:
June 21-July 5, 1941, Eastern New
England, 23
Aug. 2-3, 1941, Southern New Jer-
sey, 56

206

Field Trips (continued) :
Aug. 10, 1941, Kaiser Road, N. J.,
BY,
Sept. 28, 1941, Springdale, N. J., 22
Oct. 4, 1941, Brooklyn Botanic
Garden, 58
Oct. 18, 1941, Brooklyn Botanic
Garden, 59
Nov. 2, 1941, Delaware River to
Sunfish Pond, 118
Nov. 16, 1941, Kaiser Road, N. J.,
59
Feb. 7, 1942, Mistaire Laboratories,
Milburn, N. J., 94
Apr. 26, 1942, Bushkill Falls, 141
June 12-13, 1942, Lake Shehawken,
Ie, AY
June 13, 1942, Englewood Cliffs,
142
June 20, 1942, Branch Brook Park,
Newark, 143
June 20, 1942, Fern Garden of
W. H. Dole, 144
Oct. 4, 1942, Richmond, Staten
Island, 196
Nov. 15, 1942, Lakewood, N. J.,
197
Fiji, Botanizing in, lecture, 199
Fimbristylis, 129
Flagellaria gigantea, 126
Flora of Arizona, 9, 138; of Fukien,
190; of Indiana, 53
Forses, JAMES. Introducing Insects,
review, 50
Fraxinus nigra, 188
Frazee, V. L. 198
Frullaria Asagrayana, 141
Fuchsia arborescens, 35
Fukien, Flora of, 190
Fungi from the front lawn, 82

GaceEr, C. Stuart, 65-67

Gardenia taitensis, 127

Gentiana clausa, 188; quinquefolia,
188

Gerardia purpurea,, 188

Geum marcrophyllum,

186

186 ;

rivale,

Gity, C. L. Flowering plants and
ferns of Arizona, review, 138; and
W. H. Camp. Polypetalous forms
of Vaccinium, 168

GLEASON, H. A., 66

Glochidium ramiflorum, 126

Glyceria fluitans, 183

GoLpwalIte, RicHarp, 24

Goodyera pubescens, 184

GREENFIELD, S. Chemical inhibition
of photosynthesis, lecture, 28

Grewia crenata, 126

Grices, R. F.,°24

Guettarda speciosa, 127

GUNDERSEN, ALFRED, 59

Gunnera insignis, 37

Habenaria blephariglottis, 184; brac-
teata, 184; flava, 184

Hanp, L. E., 58, 60

Haplophyton cimicidum, 9, 10; var.
Crooksii, 9

Harpy, L., 58
Harper, R. M. Flora of Indiana, re-
view, 53

Harrts, S. K., 24

Hastincs, G. T. The plant world,
review, 18; The flower family
album, review, 51; Butterflies, re-
view, 52; Bible plants for Ameri-
can gardens, review, 114

Hedyotis foetida, 127

Helicoma, 33

Heliotropium anomalum, 127

Heracleum lanatum, 187

Hernandia Moerenhoutiana,
ovigera, 126

Hervey, ANNETTE. The use of Phy-
comyces Blakesleeanus in the assay
of thiamin in agar, lecture, 199

Heterocaryotic vigor in the Fungi,
lecture, 149

Heuchera americana, 185

Hevea, braziliensis, 110, 111

Hibiscus tiliaceus, 127; Trionum, 187

Hieracium canadense, 190; Gronovii,
190; pratense, 190

126;

207

Hires, Crara S. Field trip, Feb. 7,
1942, 94

Hopepon, ALBION, 23

HoLitincHursT, Honor M. Elected
recording secretary, 63

Hotrzorr, Mary. Field trip, June 13,
1942, 142

Horsfield, Thomas—American natu-
ralist and explorer, 1

Horsfieldia, 5

House, H. D. Clarence J. Elting and
his herbarium, 181

Houstomia purpurea, 189

Hurpary, R. A fungus disease of
Austrian pine, lecture, 28

Hybrid vigor, 102, 149

Hypericum gentianoides, 187

Aystrix patula, 183

Ilex laevitata, 187; montana, 187
Incybe eutheloides, 84; infelix, 84
Inocarpus fragiferus, 126

Iris setosa, 25

Jackson, C. F., 24

JANIGER, Oscar. The carnivorous
plants, review, 193

Jones, D. F. Chromosome relocation
and degeneration in relation to
growth and hybrid vigor, lecture,
102

Jubula pennsylvanica, 142

Juncoides, 130

Juncus, 129, 130

Jungermannia pumila, 142

Kaiser Road, New Jersey, Field trips,
57, 59

Kalmia Polifolia, 187

Karine, J. S., 65; Collecting chicle
in the American tropics, Part 1,
Soi) batt 2,09 bart. 105

Kashmir, Collecting plants in, lec-
ture, 99

Kern, F. D., 66

Korx, Laura A. More Fungi from -

the front lawn, 82
Koster, Hotuts, 56

Krigia biflora, 190; virginica, 190
KUNKEL, L. O., 66
Kyllinga, 129

Laccaria amethystina, 83; ochropur-
purea, 83

Lactuca canadensis, 190

Lake Bear Swamp, 22

Lake Shehawken, Pa., 142

Lakewood, N. J., 197

Lane, E. B. Field trip, June 20, 1942,
143

Laportea Harvey, 126

Lapsana communis, 190

Lathyrus japonicus var. glaber f.
spectabilis, 180; palustris var. myr-
tifolius, 186

Lejeunea patens, 142

Leonurus Marrubiastrum, 188

Lespedeza intermedia, 186; procum-
bens, 186; repens, 186; Stuvei, 186;
violacea, 186; virginica, 186

Leucaena glauca, 126

Leucolejeunea clypeata, 142

Levine, Micuaet. About ourselves,
Review, 86

Lewis, F. 1265

Lilium longiflorum, 99

Lily breeding, lecture, 99

Limodorum tuberosum, 184

Limonium carolinianum, 197

Lindernia dubia, 188

Lindsaya lancea, 36

Linociera pauciflora, 126

Linum virginianum, 186

Liparis lillifolia, 184

Litchfield Co., Conn., Plants of, lec-
ture, 26

Lioyp, F. E. Itinéraires botaniques
dans l’ile de Cuba, review, 191

Lobelia Canbyi, 56; Dortmanna, 189

Local names of plants, 153; Index,
166

Lonicera Caprifolium, 189

Lucuma belizensis, 75; campechiana,
75; Durlandii, 72, 75; Heyderi, 75;
salicifolia, 72, 75

208

Lunaria annua, 185

Lychnis chalcedonica, 185

Lycopodium, 36; annotinum,
tristachyum, 143

Lygodium palmatum, 95

Lysimachia vulgaris, 188

143;

Macaranga Harveyana, 126

Madia sativa, 68

Maloutia, 78

Manilkara, 77, 78, 81

Marasmius foetidus, 84

Mariscus mariscoides, 183

Marzxe, E. B. Autumn coloration,
lecture, 27; Fundamentals of plant
science, review, 90; Field trip,
Apr. 26, 1942, 142; Microscopic
anatomy in the identification of
commercial white pines, lecture, 146

McAteer, W. L. Some local names of
plants—VIII, 153

McCiintock, BAarBara. Contribution
of the nucleolus to genetic inves-
tigations, lecture, 100

McNair, J. B. Thomas Horsfield, 1

Melissa officinalis, 188

Mentha alopecuroides, 188

Menyanthes trifoliata, 188

Merritt, E. D., 66

Messerschnudia argentea, 126

Metzner, J. Observations on local
Volvocales, lecture, 26

Micromelum minutum, 126

Microspora stagnorum, 64

Mikania scandens, 189

Mimulus alatus, 188; ringens f.
roseus, 181
Mistaire Laboratories, Millburn,
N. J., 94

Mistletoe in N. J., 56

Mo.penke, H. N. Chile tarweed in
Quebec, 68; Field trip, Nov. 2,
1941, 118

Morinda citrifolia, 126; Forsteri, 126

Morus rubra, 184

Mucuna gigantea, 127

Muhlenbergia racemosa, 58, 183

Moureuy, JAMES. Field trip, Nov. 15,
1942, 197
Myrica pennsylvanica, 184

Nafaca dioica f. stellata, 179
NEARING, G. G., 58
Nephrolepis pendula, 34
Neurospora crassa, 150;
150; tetrasperma, 150
New England Tour, 23
Niue Island, Botanizing on, 121
Norton, A. H., 23
Nymphaea odorata, 185

sitophila,

Ochrosia parviflora, 127
Oenothera grandiflora, 187
Officers elected, Jan., 1942, 61
Omphalia gracillima, 84
Ophioglossum palmatum, 37; reticu-
latum, 37
Orchillium Endresti, 36
Orontium aquaticum, 184
Osiers, 11, 13
Oxalis violacea, 186
Oxycoccoides, 168-170
Oxycoccus, 168-170

Palaquim, 78

Panicum Ashet,
num, 56

Paratrophis antropophagorum, 126

Parietaria pennsylvanica, 184

Parnassia americana, 186

Paronychia canadensis, 185; fastigi-
ata, 185

Pellaea atropurpurea, 95

Pemphis acidula, 127

Peperomia, 37

PFEIFFER, NormMA E. Experiments in
connection with lily breeding, lec-
ture, 99

Phoradendron flavescens, 56

Photosynthesis, Chemical inhibition
of, lecture, 28

Phyllanthus nummulariaefolius Poir.
in the U. S., 14; corcovadensis, 15-
17; diffusus, 17; lathyroides, 15-17;
minor, 15, 17; Niruri, 15; tenel-
lus, 15-17

183 ;

Commonsia-

209

Physalis heterophylla var. nyctagi-
nea, 188; philadelphica, 188

Picea mariana, 182

Picris echoides, 190; hieracioides, 190

Pine, Fungus disease of Austrian
pine, lecture, 28; Microscopic
anatomy in the identification of
commercial white pines, lecture,
146

Pinus excelsa, 100; Lambertiana,
146; longifolia, 100; monticola,
146; Strobus, 146

Pipturus argenteus, 126

Prrone, P. P., 143

Pittosporum Brackenridget, 126

Planchonella Grayana, 127;
ensis, 126

Plantago cordata, 189

Piatt, RUTHERFORD, 62

Pleurotus hypnophilus, 84

Plumeria multiflora, 76

Podocarpus montanus, 36

Pogonia ophioglossoides, 184

Polanisia graveolens, 185

Polychytrium aggregatum, 145

Polygala Nuttallii, 56; Senega, 187

Polygonum tenue, 185

Polypodium aureum, 34, 95

Polysomatic mitosis, Prophases of,
and their relation to meiosis, lec-
ture, 200

Polystichum aculeatum plumosum,
144; andersoni, 144; Brauni, 182;
var. Purshii, 143; lonchitis, 144;
plumosum compactum, 144; wivi-
parum, 144

Pometia pinnata, 126

Potamogeton bupleuroides, 26; pecti-
natus, 182

Potato chips, lecture, 97

Potentilla Anserina, 186; palustris,
186

Premna taitensis, 127

Proceedings of the Club:
Meeting of

Oex 19, WO, 25
Nov. 3, 1941, 26

SAMO-

Nov. 19, 1941, 27
Dec. 2, 1941, 29
Dec: 17, M041" 729
Jan. 6, 1942, 61
Jan. 21, 1942, 62
Feb. 3, 1942, 97
Feb. 18, 1942, 98
Mar. 3, 1942, 99
Mar. 18, 1942, 100
Apr. 7, 1942, 101
Apr. 15, 1942, 145
May 5, 1942, 146
May 20, 1942, 147
Oct. 6, 1942, 148
Oct. 21, 1942, 149
Nov. 2, 1942, 198
Nov. 18, 1942, 198
Dec. 1, 1942, 199
Dec. 16, 1942, 200
Procris pedunculata, 126
Proserpinaca palustris, 187
Prunus cuneata, 58
Psalliota abruptibulba, 85
Pseudolmedia oxyphyllaria, 75
Psidium Guajava, 126
Psilocybe foenescti, 85
Psychotria insularum, 126
Pyrola secunda, 187

Quercus prinoides, 60; X Schuettet,
184

Ranunculus ambigens, 185; delphim-
folius, 23; flabellaris, 185; micran-
thus, 185 Z

Rauwolfia Spectabilis, 77, 81

Reep, G. M., 67

Rhipidopteris peltata, 35

Rhododendron canadense, 58,
maximum, 187

Rhus taitensis, 126

Ribes odoratum, 186

Richmond, S. I., 196

Rickett, H. W. The names of
Cornus, 11; An Herbal, review,
91; Cornus again, 131; John Tor-
rey, review, 135

RIpvLe, Oscar, 67

187;

210

Rozssins, W. J., 66, 67; Vitamins
and growth of plants, lecture, 30

ROUSSEAU, JACQUES, 25

Rudbeckia speciosa, 189

Rusk, Hester M. Field trip, Oct. 4,
1942, 196

Russula variata, 82

Rynchospora, 129; alba, 183; knie-
Skernii, 56

Sagittaria latifolia f.
183; subulata, 182

Salvia coccinea, 127

Samolus parviflorus, 188

Sanguisorba canadensis, 186; minor,
186

Sapium, 78

Sapodilla, 38, 39, 69, 105-111

Saponaria Vaccaria, 185

Sarracenia purpurea, 185

Saxifraga pennsylvanica, 185

Scabiosa arvensis, 143

Scaevola frutescens, 126

Scleria, 129

Scumipt, Mary. Demonstration, 151

Scirpus, 129; hudsonianus, 183

Scleroderma aurantium, 84; cepa,
84; lycoperdoides, 84; tenerum, 84

Scutty, F. J. Sedges and rushes of
Hot Springs National Park, 129

Scutellaria parvula, 188

Sedges and rushes of Hot Springs
National Park, 129

SerFriz, W. Recent advances in the
study of protoplasm, lecture, 29

Senecio Cooperi, 35; obovatus, 190

Seventy-fifth anniversary célebra-
tion of the Club, Program, 65

Shehawken, Lake, 142

SHUEEa Ga rl) 67

Sideroxylon, 78; amygdalinum, 72,
74; Gaumeri, 72, 74; Meyeri, 72,
74

Sigma Delta Epsilon fellowship, 151

Silene Armeria, 185; stellata, 185

Sinnott, E. W., 65

SmatL, J. A., 68; Field trips of
June 21-July 5, 1941, 23; Aug. 2-3,

diversifolia,

1941, 56; Aug. 10, 1941, 57; Sept.
28, 1941, 22; Oct. 4, 1941; 58;
Oct. 18, 1941, 59; Nov. 16, 1941,
59

Smilacina racemosa, seed dormancy,
64

Smilax laurifolia, 56; Walteri, 56

Solanum rostratum, 188

Solidago neglecta, 189; odora, 189;
patula, 189; ulmifolia, 189

Sorbus americana, 186

Southern New Jersey field trip, 56

Spargamum americanum, 182

Spartina cynosuroides, 197

Specularia perfoliata, 189

Spinacia oleracea, Polysomatic mito-
sis in, 200

Spiranthes plantaginea, 184

Sporodinia grandis, 86

Springdale, N. J., 22

STEINMETZ, F. H., 24

Stellaria borealis, 185

Stemmadenia Donell-Smithi, 75

Stenophyllus capillaris, 183

Stewart, R. R. Collecting plants in
Kashmir, lecture, 99; Flora of
Fukien, review, 190

Stout, A. B. Standardized plant
names, review, 132

Svenson, H. K., 66; Vegetation of
western South America, lecture,
198

Svida, 11, 13

Tabernemontana, 76

Tarenna sambucina, 126

Taxodium distichum, 57

Taytor, Norman, 151

Terminalia Catappa, 127

Thespesia populnea, 127

Thevetia nitida, 76

Tuomson, J. W., resigned as record-
ing secretary, 63

Tuornton, N. C. The mystery of
the potato chip, lecture, 97

Timonius polygamus, 127

Torrey, John, 135

Trema orientalis, 126

211

Trichomanes lucens, 34

Triodia eragrostoides, 9, 10; flava,
183

Triumfetta procumbens, 127

Trollius laxus, 185

Utricularia cornuta, 189; juncea, 189

Vaccinium, Polypetalous forms of,
168; altomontanum, 170; arkan-
sanum, 170; atrococcum, 170, 172;
brittonu, 172; corymbosum, 170;
macrocarpon, 187; missourtense,
169, 170; pallidum, 169; subcorda-
tum, 169; tallapusae, 170; torrey-
anum, 169-171; vacillans, 169, 170;
var. crinitum, 170; viride, 169, 170

Verbena stricta, 188

Vernoma noveboracensis, 189

Veronica <Anagallis-aquatica, 188;
longifolia, 188

Viburnum prunifolium, 189

Vicia Cracca, 186

Viola canadensis, 187; lanceolata,
187

Virea autumnalis, 190
Vitamins and growth of plants, lec-
ture, 30

Vitis Labrusca, 187

Volvocales, Local, lecture, 26

Volvox aureus, 26; globator, 26;
weismannia, 26

Waldsteinia fragarioides, 186

Wetmore, R. H., 65

Wuatey, W. Gornon. Practical plant
anatomy, review, 88; Methods of
plant breeding, review, 137

Wirkus, ELEANOR and C. A. BERGER.
Polysomatic mitosis, lecture, 200

Witte, C. P., 143

Woodsia obtusa, 182

Xylaria polymorpha, 82
Xyris flexuosa, 184

Yuncxer, T. G. Botanizing on Niue
Island, 121

Zamia Skinneri, 36

Zanthoxylum americanum f. impu-
niens, 180

ZIMMERMAN, P. W., 66; Elected to
Council, 97

Zizia cordata, 187

Zosterella dubia, 184

by ah iene «

THE TORREY BOTANICAL CLUB

William J. Robbins
C. Stuart Gager
John S. Karling
Fred J. Seaver

1941-1943
John H. Barnhart
R. C. Benedict
Helen M. Trelease
P. W. Zimmerman

Clarence Lewis, Chairman
J. Ashton Allis
Caroline C. Haynes

Edwin M. Matzke, Chairman (ex officio)

Charles A. Berger
Arthur H. Graves
Honor M. Hollinghurst

Edward J. Alexander
Vernon L. Frazee
Eleanor Friend
Alfred Gundersen
Robert Hagelstein

William J. Bonisteel
Herbert M. Denslow
James Edwards

Ferns and Fern Allies:

Council for 1943

Ex officio Members
Lela V. Barton

Edwin B. Matzke
Honor M. Hollinghurst
W. Gordon Whaley

Elected Members

1942-1944
J. M. Arthur
W. J. Bonisteel
Arthur H. Graves
Sam F. Trelease

Committees for 1943

ENDOWMENT COMMITTEE

Harold W. Rickett
Michael Levine
John A. Small
Bernard O. Dodge

1943-1945
Charles A. Berger ,
Clyde Chandler
Albert E. Hitchcock
Roger P. Wodehouse

Henry de la Montague
Helen M. Trelease

ProGRAM COMMITTEE

A. B. Stout

W. Gordon Whaley
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FIELD COMMITTEE
Joun A. Smaty, Chairman

Inez M. Haring
Michael Levine
H. N. Moldenke
James Murphy
G. G. Nearing

LocaL Frora COMMITTEE
Epwin B. MatzKeE, Chairman

Dolores Fay
John M. Fogg, Jr.
H. Allan Gleason

Cryptogams

Mosses: E. B. Bartram
Liverworts: A. W. Evans, E. B. Matzke
Freshwater Algae: H. C. Bold

Marine Algae:

J. J. Copeland

S. Karling

Fungi: A. H. Graves, J.
Lichens: J. W. Thomson, Jr.
Myxomycetes: R. Hagelstein

PUBLICATIONS EXCHANGE COMMITTEE

Edwin B. Fatzke, Chairman (ex

officio) Amy L. Hepburn

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Harold W. Rickett
Hester M. Rusk
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R. C. Benedict, W. Herbert Dole, N. E. Pfeiffer

Lazella Schwarten

OTHER PUBLICATIONS
OF THE

TORREY BOTANICAL CLUB

(1) BULLETIN

In addition to papers giving the results of research, each issue
contains the INDEX TO AMERICAN BoTANICAL LITERATURE—a very
comprehensive bibliography of current publications in American
botany. Many workers find this an extremely valuable feature of the
BULLETIN.

(2) MEMOIRS

Volume 18, no. 1, 108 pages, 1931, price $2.00. Volume 18, no.
2, 220 pages, 1932, price $4.00. Volume 18 complete, price $5.00.

Volume 19, no. 1, 92 pages, 1937, price $1.50. Volume 19, no.
2, 178 pages, 1938, price $2.00.

(3) INDEX TO AMERICAN BOTANICAL
LITERATURE

TORKEYA

A Bi-MonTHLY JOURNAL oF BoTanicaL Notes AND NEws

John Torrey, 1796-1873

EDITED FOR

THE TORREY BOTANICAL CLUB
BY

BUNIROIUID) Ist, CILIUM

VOLUME 43

New York

1943

aed
Wey 4

+r Oe

BOTANICAL

Volume 43 July 1943 GARDEN Number 1

TORREYA

EDITED FOR
THE TORREY BOTANICAL CLUB

HAROLD H. CLUM

John Torrey, 1796-1873

CONTENTS

THE SEVENTY-FIFTH ANNIVERSARY CELEBRATION OF THE TORREY
BOTANICAL CLUB, JUNE 22-27, 1942

Introduction
Haphazard as a Factor in the Production of Tetrakaidecahedra..FRrEDERIC T. Lewis 4
The Evolution and Determination of Sexual Characters

in the Angiosperm Sporophyte.................2+.e0ece00 CHARLES FE. ALLEN 6
Leaf-stem Relationships in the Vascular Plants............... RatpH H. Wetmore 16
Cell Division as a Problem of Pattern in Plant Development..EpMUND W. Sinnott 29
Contributions of the Torrey Botanical Club to the Development of

PAN ONOMLY, 252 hes chee PuSine aon slays cle SR aM DDS aa Vere to Ceara Ue Anage H. A. Gieason 35
Modern Taxonomy and Its Relation to Geography............ Henry K. Svenson 44
Some Economic Aspects of Taxonomy..................02:02002%: E. D. Merritt 50
The Importaunce of Taxonomic Studies of the Fungi.............. Frank D, Kern 65

Activities of the Club, January to May 1943. ........... 0. ccc cece ec cee ees c cee eseenee 78

PUBLISHED FOR THE CLUB

By THE FREE PRESS Printing CoMPANY
187 CoLLEGE STREET, BuRLINGTON, VERMONT

Entered as second class matter at the post office at Burlington, Vermont,
October 14, 1939, under the Act of March 3, 1879

THE TORREY BOTANICAL CLUB
OFFICERS FOR 1943

President: WiLttAM J. Roppins

Ist Vice-President: Frep J. SEAVER Recording Secretary: Honor M. Hot-

2nd Vice-President: LELA V. BARTON LINGHURST

Corresponding Secretary: EDw1n B. MATzKE Treasurer: W. GorDoN WHALEY
Editor: Harotp W. RICKETT

Associate Editors:

Irvinc W. BaAILey ADRIANCE S. FOSTER
EpwaArp W. Berry Henry A. GLEASON
STANLEY A.,CAIN ARTHUR H. GRAVES
M. A. CHRYSLER ' JoHNn W. SHIVE
Harotp H. CLum R. P. WoDEHOUSE

MicHAEL LEVINE
Business Manager: MicHAEL LEVINE Bibliographer: Mrs. LazeELLa SCHWARTEN
Delegate to the Council, N. VY. Academy of Sciences: BERNARD O. DopcE

Representatives on the Council of the American Association for the
Advancement of Science

Joun H. BARNHARDT ALBERT F. BLAKESLEE

Representative on the Board of Managers of the N. Y. Botanical Garden:
Henry A. GLEASON

MEMBERSHIP IN THE TORREY BOTANICAL CLUB

All persons interested in botany are invited to join the club. There are four classes
of membership: Sustaining, at $15.00 a year; Life, at $100.00; Annual, at $5.00 a year
and Associate, at $2.00 a year. The privileges of members, except Associate, are: (a) To
attend all meetings of the club and to take part in the business, and (b) to receive its
publications. Associate members have the privilege of attending meetings, field trips
and of receiving the Schedule of the Field Trips and the Bulletin of the New York

Academy of Sciences.
TORREYA

TorREYA was established in 1901 as a monthly publication of the Torrey Botanical Club
for shorter papers and interesting notes on the local flora range of the Club. It also
contains the proceedings of the Club, reports of field trips, and some book reviews and
news notes. The Council of the Torrey Botanical Club has decided to devote volume 43
of TorreEyYA, 1943, to the publication of the papers presented in June 1942 at the 75th Anni-
versary Celebration of the Club, and to the Proceedings of the Club. This volume will
be published. in two numbers.

ToRREYA is furnished to subscribers in the United States and Canada for one dollar
per year (January-December): single copies thirty cents. To subscribers elsewhere,
twenty-five cents extra, or the equivalent thereof. Postal or express money orders, drafts,
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Of the annual membership dues of the Torrey Botanical Club, $.50 is for a year’s
subscription to ToRREYA.

ToRREYA is edited for the Torrey Botanical Club by

HAROLD H. CLUM
HUNTER COLLEGE, 695 Park AVENUE
New York, N. Y.

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Vor. 43 Jury 1943 No. 1

Introduction

The Torrey Botanical Club is the oldest botanical society in America, and
ever since its founding, its members have been active in all botanical move-
ments, such as the discussions of rules of nomenclature, the establishment oi
the Botanical Section of the American Association for the Advancement of
Science, and of other botanical organizations ; and at the Semicentennial Cele-
bration of the Club in 1917, preliminary discussions were held which have since
led to the establishment of “Botanical Abstracts,’ now “Biological Abstracts.”
With this record of botanical achievement it seemed fitting that a Seventy-fifth
Anniversary Celebration should be held. In the fall of 1941 Dr. J. S. Karling,
then President of the Club, appointed a large committee to discuss the pos-
sibilities of such a celebration. It was decided to hold the celebration in New
York in June 1942, independently of meetings of the American Association ot
the Advancement of Science and of other botanical societies. However, all
botanists were invited to participate. It was also decided to hold four sessions.
at which papers would be presented, in four different local institutions; te
leave the afternoons free for recreation and inspection of these institutions:
and to provide some evening entertainment and a field trip. Smaller com-
mittees were appointed to work out the details of securing speakers, and of
arranging for the accommodation of visitors and delegates. As events worked
out, the summer scientific meetings at Ann Arbor, Michigan, were canceled
and the Botanical Society of America joined in the celebration of the Club
in lieu of a separate summer meeting in the East.

Registration began Monday morning, June 22, 1942, at Columbia Univer-
sity. At 2:00 p.m. the first session was called to order by Dr. E. B. Matzke.
Dr. Karling gave an interesting review of the development of the Department
of Botany at Columbia from the early days before the University occupied
its present site, and told of the establishment of the Elgin Botanical Garden
as an aid to the teaching of botany. Then followed the papers by Drs. F. T.
Lewis, C. E. Allen, R. H. Wetmore, and E. W. Sinnott. These are presented
here, although the papers of Dr. Lewis and Dr. Sinnott are in somewhat
abbreviated form.

After the formal program the visitors resorted to the Low Memorial
Library for tea, and to examine a display of books and reports by John Torrey,
and photographs of former botanists at Columbia and of others associated
with the Torrey Botanical Club in earlier days. In the evening the anniversary

Torreya for July (Vol. 43, 1-85) was issued August 27, 1943

INTRODUCTION 3

banquet at the Men’s Faculty Club was well attended and proved to be an
enjoyable occasion with Dr. C. Stuart Gager, President of the Club, presiding.
Excerpts were read from many letters of felicitation from various organiza-
tions and from individual botanists who could not come.

On Tuesday morning the meeting was held at the New York Botanical
Garden with Dr. William J. Robbins presiding. Dr. Robbins first gave a very
interesting account of the history of the Botanical Garden, illustrated with a
number of slides showing the activities of the Torrey Botanical Club in the
establishment of the Garden, and in the erection of the museum building and
greenhouses. Following this talk four papers on different aspects of taxonomy
were given by Drs. H. A. Gleason, H. K. Svenson, E. D. Merrill, and
F. D. Kern. These make up the last half of this number of Torreya. At the
end of the program the accompanying picture of the group was taken on the
steps of the museum building. The weather still seemed too threatening for
lunch to be served out-of-doors, and arrangements were made for it on the
main floor of the museum building. Fortunately, however, it cleared sufficiently
for the inspection of the gardens in the afternoon.

On Wednesday, June 24th, the program was continued at the Boyce
Thompson Institute for Plant Research. Here Dr. P. W. Zimmerman presided
and Dr. William Crocker told of the establishment and growth of the Institute
during the past twenty years. Three papers on growth problems were pre-
sented by Drs. L. O. Kunkel, P. W. Zimmerman, and O. Riddle. Following
the program the Institute served a very nice luncheon; and then dividing the
visitors into small groups, the staff of the Institute conducted everyone through
the building and greenhouses on exceedingly well organized tours with a
minimum of congestion or confusion. As reference was made in the last para-
graph to threatening weather, and rain will be mentioned again toward the
end of the volume, it is perhaps excusable to state that this was a perfect June
day, and the rose arbor at the Institute was at its height of bloom.

In the evening Dr. William J. Robbins gave a lecture on vitamins at the
American Museum of Natural History.

On Thursday the group met at the Brooklyn Botanic Garden. Dr. C. Stuart
Gager presided and told of building the Botanic Garden, which has many
phases of activity, is composed of a number of diverse unit gardens, and serves
the public in many ways, on an originally unattractive piece of waste land.
Four papers were given in the formal program by Drs. G. H. Shull, S. A. Cain,
G. M. Reed, and A. F. Blakeslee. Luncheon was served in the Brooklyn
Museum, and this was followed by an inspection of the gardens.

The papers presented on Wednesday and Thursday, and an account of
the field trip of Friday and Saturday, will be published in the second number
of TorREYA. EG

Vor. 43 190 KORE WA Jury 1943

Haphazard as a Factor in the Production of Tetrakaidecahedra*

FrReDERIC T. Lewis

This paper, more fully presented than was possible in oral delivery and
with added reference to subsequent publications, has been published in the
AMERICAN JOURNAL OF Botany, 30: 74-81. Jan., 1943. There it is entitled
“A Geometric Accounting for Diverse Shapes of 14+-hedral Cells: the Transi-
tion from Dodecahedra to Tetrakaidecahedra.” A summary of the discourse
follows :

The study of cell shapes in compact parenchyma, or in similar unspecialized
aggregates, has led to a series of surprises. (1) Such cells, instead of being
rhombic dodecahedral products of surface tension, in reality have an average
of between 13.5 and 14 facets,—usually close to 14. (2) The cells, though
having an average of 14 facets, very rarely present the 14~hedral shapes
deduced by Lord Kelvin as dividing space into uniform bodies of minimal
surface. Even irregular or distorted approximations of those shapes, with 8
irregularly hexagonal facets and 6 quadrilaterals that are far from true squares,
occur in less than 1 per cent. of the cells studied. (3) Compressed solids,
such as shot of a given size, no longer controlled by surface tension, assume |
the same irregular cell-like shapes with the same average of close to 14 facets |
(Marvin). (4) Aggregations of soap bubbles of as nearly uniform size as
they can be made, responding to surface tension, and free to glide over one
another, do not assume the Kelvin shapes. With an average of 14 facets, they |
present a variety of cell-like forms (Matzke).

Confronted with this situation, the aggregation of geometrically perfect |
rhombic dodecahedra was considered anew. At six corners of each rhombic
dodecahedron, when surrounded by others like it, six polyhedra meet at a
mathematical point. Let two of them deviate a hair from meeting the other four |
at a point, and let the deviations throughout the mass occur in all directions |
at random, and the aggregation of rhombic dodecahedra becomes an assem- |
blage of irregular shapes with an average of 14 facets. The shapes range from |
12— to 18-hedra, and have an abundance of pentagonal facets. When all edges |
are more or less of the same length, these irregular polyhedra present many of |
the forms common to cells, bubbles in foam, and compressed shot. The average |
of close to 14-facets in a disarranged space-filling mass of bodies of similar
size thus appears inevitable. The occasional occurrence of five or six cells |
chancing to meet at a point, or of four meeting along a line, would slightly |
reduce the average.

* Read at the 75th Anniversary Celebration of the Torrey Botanical Club at Columbia |
University, Monday, June 22, 1942.

LEWIS: PRODUCTION OF TETRAKAIDECAHEDRA 5

Cell division extends the difference in the number of facets both above
and below the afore-mentioned range of 12 to 18. Yet if the average plane
of division is hexagonal (which would be expected when division bisects a
cell rather than cuts off a corner) it will not affect the average of 14 facets.
It causes a diversity in cell size, incompatible with a full realization of the
Kelvin pattern. Yet if division occurs in a prevailing plane, it can orient
the cells, and orientation makes possible an approach to Kelvin’s orthic 14—
hedron, which approximation is indubitably present in the oriented pith of
Eupatorium and in similar tissue.

We conclude, therefore, that an average of 14 facets can be due to chance,
or to tension, or a combination of both. The mathematical solution of the
problem of dividing space into uniform bodies of least surface area and of
maximum stability has been solved by Lord Kelvin’s minimal 14—hedron (or
its close approach,—his orthic 14-hedron). Since such diverse forms as the
stellate 12—-rayed cells of Juncus, and the prosenchymal tracheids of the pine
with from 18 to 22 facets apiece, are accountable as derivatives of the Kelvin
14-hedron, as well as all the forms in cork and pith, it may properly be
regarded as the typical shape of cells in masses. There is no rival uniform
pattern. But it is only through absolute uniformity in size, precision in align-
ment, and the dominance of surface tension (3 factors at least) that a foam
of minimal 14-hedra may be expected. These conditions have apparently
not yet been realized in any cells or any froth. The typical shape thus remains
a mathematical abstraction, whereas the actual shapes are coming to be well
understood, and haphazard is a factor.

Harvarp Mepicat SCHOOL
Boston, MASSACHUSETTS

Vor. 43 ORR RA NON JuLy 1943”

The Evolution and Determination of Sexual Characters in the
Angiosperm Sporophyte*

CuHarLes E. ALLEN

One result of genetic study which bears definitely upon evolutionary
theory is the demonstration that the determination of an apparently simple
character depends upon the activity of many genes. It is indeed suggested
that the interaction of all the genes of an organism may be essential to the
appearance of any character; but for the present this broader conception
remains in the realm of speculation.

Another contribution from the same source is the demonstration that
similar or apparently identical phenotypes may be determined by diverse
genic complexes. It follows that very different gene mutations in distinct lines
of descent may result in the appearance of similar characters—a fact which
in another aspect students of phylogeny have long stated in terms of parallel
or convergent evolution.

Turning to a special class of characters, it is evident that sexual dif-
ferentiation has arisen independently in many different plant and animal
lines. There is no reason for assuming that the changes in the genetic
mechanism which resulted in this differentiation were identical, or even
closely similar, in diverse lines.

To this consideration is to be added that, after the first step in sexual
differentiation, additional mutations occurred, independent and varying in
different lines. These later steps resulted in differentiation between the organs
in which gametes are produced; in differentiation of individual gamete-
producing plants or animals as respectively female and male; and, in certain

pteridophytes and in all seed plants, in a backward extension of sexual |

differentiation to involve structures of the parental spore-bearing generation.
A priori, then, it is not to be expected that the genetic mechanisms which

determine sexual potentialities or which influence sex-expression should be |
the same in different groups of organisms. Yet it is characteristic of discus- |

sions in this field that unitary theories of “sex-determination” have been
developed; each based upon phenomena observed in one or in a few related

species, but each seeking to apply one mechanism to all groups of sexually |

differentiated organisms. There is, to be sure, one set of facts which may

seem to support this conception of uniformity: namely, the occurrence in |

widely separated phyla of apparently similar bodies—the sex chromosomes—

which are a part of the genetic mechanism whose nature is being sought. |

* Read at the 75th Anniversary Celebration of the Torrey Botanical Club at Columbia |

University, Monday, June 22, 1942.

att

ALLEN: EVOLUTION OF SEXUAL CHARACTERS

™N

But it would not be surprising to discover that this type of similarity presents
an additional instance of parallel evolution.

The history of angiosperms begins at a level at which the greatly reduced
members of the gamete-bearing generation had long been sharply separated
as female and male individuals. Sexual differentiation also had been projected
backward to effect a distinction in the parental generation between female
and male spore-bearing structures. These structures—macro- and micro-
sporangia—were now borne upon or within likewise sexually differentiated
organs later to be designated pistils and stamens.

Less confident must be any statement as to the distribution of pistils and
stamens in primitive angiosperms. Three conditions are conceivable. Either
the original angiosperm flower was bisexual (bisporangiate), the plant bearing
it being hermaphroditic; or there were separate pistillate and staminate
flowers, borne either on the same plant (a condition of monoecism) or on
distinct plants (a condition of dioecism).

Attempts to choose between these possibilities were based first upon
comparative morphology; then, as fossil evidence accumulated, the assistance
of paleobotany was sought. The latter source has as yet contributed little
to the problem here involved. It has shown that the equivalent of a bisporan-
giate flower was developed by Cretaceous times in the Bennettitales ; and that
the equivalent of a unisporangiate flower was present in the Caytoniales as
early as the Triassic. But it is agreed that neither Bennettitales nor Cay-
toniales were ancestral to modern angiosperms. Probably the great majority
of those who have discussed the question have concluded that primitive angio-
sperms had bisexual flowers. But unanimity upon this point is not reached ;
and the possibility of a polyphyletic origin, some lines starting with herma-
phroditism, others with monoecism or dioecism, is not wholly excluded.

Since the sharp distinction between female and male gametophytes was
established at a pre-angiosperm level, a discussion of the evolution of sexual
characters in angiosperms can deal only with developments within the spore-
bearing generation. It may be asked, first, what if any genetic evidence is
there as to the type of distribution of sexual structures in primitive angio-
sperms? Second, what appears to have been the most probable course or
courses of evolution of sexual characters since the dawn of angiosperm history ?

Two general sets of facts, long recognized and both to be referred to
later, suggest the derivation of unisexual from bisexual flowers. One of these
concerns the presence in the majority of monoecious and dioecious species
of pistil-rudiments in staminate flowers and of stamen-rudiments in pistillate
flowers. The stage to which these rudimentary structures develop varies from
that of a small hump of undifferentiated tissue to that of the reflexed stamens
of the functionally female flowers of the grape, which produce non-viable or

[e,0)

AOR YA

rarely viable pollen. The appearance of such variably developed but func-
tionless or chiefly functionless organs is difficult to explain save by reduction
in the course of descent from a hermaphroditic ancestor. A different con-
ception may be based upon those species whose unisexual flowers show no
trace of organs of the opposite sex. When, however, such plants belong to,
or are obviously closely related to, families among whose members are some
with bisexual flowers ocr with unisexual flowers containing staminodia or
pistillodia, the obvious explanation of unisexuality is still that of descent
by reduction from a bisexual condition. There remain the relatively tew
species that supply no indication, through either structure or relationship,
of such descent. It was this condition in Casuarina which made it, in Wett-
stein’s phylogenetic scheme (19), the starting-point for angiosperms.

The other set of facts with a similar bearing is the variability of sexual
conditions in those angiosperms whose flowers are typically unisexual. In
many monoecious and dioecious species, bisexual flowers now and then appear.
Even more frequently, staminate replace pistillate flowers and vice versa;
flowers of either sex appearing on the dioecious plant or on the part of the
monoecious plant which regularly bears flowers of the opposite sex. Whatever
its explanation, such lability of sex-expression in the sporophyte contrasts
sharply with the rigid separation of sexual characters in the gametophyte.
Comparable lability seems to characterize gyno- and andromonoecious, gyno-
and androdioecious species.

In this connection, too, cases may be found which could be thought to
point in an opposite direction. Among the dioecious species that have been
extensively studied, three (two of Lychnis and one of bryonia) present a very
sharp sex-separation. Doubtless, when other less well-known species are
studied, similar instances will be found. But in Bryonia dioica staminate
flowers, in Lychnis dioica and L. alba both staminate and pistillate flowers,
contain rudiments of organs of the opposite sex. The change in sex-expression
in pistillate flowers or Lychnis under the influence of the anther smut is
well known, although human ingenuity has yet found no means of accom-
plishing a like result. No modification of sex-expression is known to have
been induced in Bryonia dioica, though there is one old report of a female
plant bearing some bisexual flowers.

The common variability of the unisexual condition is among the facts
which long ago led to the conclusion that in all angiosperms genotypic bases
are present for both femaleness and maleness. No reason has appeared to
question this conclusion; indeed, all later-adduced evidence has but served
to confirm it. Correns (2) postulated for dioecious species an additional gene
or gene-complex for sex tendencies superposed upon those representing sex
potentialities. This formulation, recognizing a certain degree of genotypic

ALLEN: EVOLUTION OF SEXUAL CHARACTERS 9

complexity, is still, so far as it goes, valid as a formal statement of the case.
As will appear, it is now evident that the story is even more complicated than
Correns’ statement would imply.

A first suggestion of the complexity to be expected appears in the fact
that in dioecious angiosperms as in metazoa there are two general types of
genotypic influence upon sex. In one, the more common type in both groups,
the male is heterozygous, the female homozygous, for sex-tendency factors
as well as for sex chromosomes. In the other, represented by strawberries and
possibly by members of a few other genera, the male 1s homozygous, the female
heterozygous. The difference is explicable by descent from hermaphroditic
ancestors, different mutations in which have led to opposite results. It adds
to the improbability of an assumption of the primitiveness of dioecism.

A type of mutation observed in a considerable number of hermaphroditic
species involves a stoppage at some stage in the development of stamens (or
their complete failure to develop), with the result either that no pollen is
produced or, if produced, it is nearly or quite functionless. “Male-sterile”
mutations of this general nature have been studied, for example, in the sweet
pea, shepherd’s purse, sorghum, Oenothera, onion, tomato, potato, barley (16).
In these and in other plants, the condition in question seems to be due to a
mutated gene (or to at least two genes in the tomato), the mutation being
always or nearly always recessive. Mutations of a somewhat different sort
bring about a replacement of stamens by petals or petaloid structures. It is
clear that mutations of both types have occurred on a large scale in the past ;
witness the frequent occurrence, previously noted, of staminodia or stamen
rudiments replacing some of the stamens in bisexual flowers or all the stamens
in flowers which are now unisexual. Notable are the partial petaloid trans-
formation of the last-remaining stamen of Canna; the often-observed occur-
rence of doubleness in consequence of a transformation of stamens; and the
evidence from the morphological side that petals in many cases represent
transformed and sterilized stamens. Mutations tending toward male sterility
occur likewise in monoecious species. In maize, the most studied genetically
of all plants, at least 27 distinct mutations of this general nature have been
observed (7); 20 classed as ‘“‘male-sterile,’ 5 as “tassel ear,’ one each of
“antherless” and “pollen lethal.’ These 27 mutations involve as many distinct
gene loci; all but two are recessive.

Such mutations in the direction of male sterility might be described as
tending toward femaleness. Those of another type, known for example in
Silene, Cheiranthus, and Papaver, in which stamens are replaced by carpels,
may be similarly classed.

Comparable with the mutations which result in or tend toward male
sterility are those leading toward female sterility. The striking fact shown

10 TORSRIE Yo

by a review of the literature is that mutations of this type appear, in both
hermaphroditic and monoecious species, to be far less frequent than are
those leading toward male sterility. Female sterility, pistils being more or
less aborted, seems to be recessive in mutants of Phlewm pratense, Antirrhi-
num, rice, and raspberries. In calycanthema forms of Campanula and of
Rhododendron, the mutant condition (pistils developed but sterile) is domi-
nant. In Geranium, pistils functional as such but showing structural transitions
toward the staminate character have appeared in interracial crosses. The
behavior of the character in back-crosses, while not entirely clear, suggests
a Mendelian segregation. In a cross between species of Geum, the results of
further matings are likewise not clear-cut. It is possible that in these crosses,
as apparently in a few interspecific crosses which have resulted in male
sterility, cytoplasmic influences are involved.

In the same list of mutations in maize which shows 27 genes involved in
male sterility, only 6 mutations leading toward female sterility are reported:
two “anther ear,” 2 “barren stalk,’”’ 1 “lethal ovule,” 1 “silkless.”

To the story of observed mutations in this direction must be added the
known cases of pistillodia, which represent the result of past mutations;
possibly the reduction in number of ovules in certain lines; and the relatively
few cases of doubleness which have involved the transformation of pistils
as well as of stamens into petals.

To be mentioned also are a few known mutations which, like two observed
in intervarietal crosses in Oryza (12), tend simultaneously toward both male
and female sterility. Obviously mutations of this class can hardly have played

a direct part in evolution. In general they seem to result in monstrosities |

which, even apart from the accompanying sterility, would probably not be
favored by selection. Chromosomal changes may be involved.

It is not yet clear why mutations toward male sterility are much more
frequent than those toward female sterility. This difference holds not only
for observed mutations. As to the past, it is evident that petals in a large
proportion of instances represent sterilized stamens; only rarely can they
be considered sterilized carpels.

The mutations thus far cited involve changes in the general direction
from hermaphroditism toward dioecism. As already mentioned, very many
variations (to be distinguished from mutations) occur in the opposite direc-
tion—involving the appearance in monoecious or dioecious species of bisexual
flowers or, in dioecious species, of both pistillate and staminate (and some-
times bisexual) flowers on the same plant. Such variations are in large
measure shown to be reactions to environmental conditions. They may be
considered expressions of genotypic possibilities present from a remote

——

eos

ALLEN: EVOLUTION OF SEXUAL CHARACTERS 11

ancestry, certain of which have been inhibited, but not completely suppressed,
by mutations like those previously mentioned.

But apart from variability of this common type, now and then a demon-
strable mutation occurs in the direction from dioecism or monoecism toward
hermaphroditism. Such mutations are known in dioecious species of Lychnis,
Salix, Silene, Vitis, and Fragaria; in Salix and Silene they have occurred
in the offspring of interspecific crosses. One recessive mutation resulting
in bisexual flowers is known in maize (7). Most monoecious or herma-
phroditic strains derived from dioecious species and subjected to genetic
experiment have behaved as though they were mutated males; a very few
have seemed to be mutated females. In Lychnis both mutated males and
mutated females have been recognized cytologically. Those hermaphrodites
(the term 1s often somewhat loosely used) which appear to be mutated males
in general behave in breeding like males; that is, their progeny shows them
to be heterozygous for a sex-tendency gene. In this respect they differ from
regularly hermaphroditic species, which of course transmit hermaphroditism
uniformly to all their progeny. This genotypic difference, as Correns pointed
out, justifies the description of the appearance of hermaphroditism in a
dioecious or monoecious species as a case of “backward evolution.” The implica-
tion is that in a dioecious species derived from a primitively hermaphroditic
one a mutation has produced a reversion to the phenotypically original char-
acter—although this change is not due to a reverse mutation of a previously
mutated gene.

Not always readily distinguishable from these variations and mutations
are the reported cases, in species classed as dioecious, of strains which regu-
larly vary in degree of sex-separation. In Urtica cannabina, Spinacia, and
Mercurialis, for example, plants shown to be genetically distinct occur which
are monoecious or hermaphroditic. In other instances differing degrees of
sex-separation are manifested by different strains. Comparable but not fully
elucidated cases are presented by gynodioecious species. In the absence of
direct evidence as to their origin, these diverse conditions are capable oi
explanation either as steps in an evolutionary sequence leading toward
dioecism, individuals showing intermediate conditions not yet having been
eliminated; or as evidences of mutation in the reverse direction comparable
with the cases studied in Lychnis.

Mutations, then, may and do occur both in the general direction from
hermaphroditism toward dioecism and in that from dioecism toward herma-
phroditism. Those of the latter class are much the less frequent, and the
best known of them lead to a hermaphroditism which is not genetically like
the hermaphroditism which may be considered primitive. It is evident, too,
that mutations away from hermaphroditism have been numerous in the past

12 WOR Ran YY AL

and that, as shown by the persistence of rudimentary structures, very many
of them have become fixed as part of the specific genotype.

The conclusion indicated by genetic evidence hence agrees with that most
strongly suggested by morphological study; the general tendency in angio-
sperm evolution has been from a primitive hermaphroditism toward dioecism.
In many species various intermediate stages have been reached; in something
less than 5000 according to available counts (20), the final step to dioecism
has been taken.

The mutations that have been chiefly concerned in the evolution of sexual
conditions in angiosperms have involved a diminution or loss of the power
of spore-production; commonly also a loss or reduction of the organs con-
cerned. The mutations of this nature which are appearing at present are with
rare exceptions recessive. It is reasonable to assume that similar mutations
in the past history of angiosperms have, at their origin, likewise been chiefly
recessive. Those mutations which have played the major role in floral evolu-
tion agree, therefore, in two respects—in involving a loss or diminution of
potentialities and in being originally recessive—with the general run of
observed mutations in all organisms. So far, then, as concerns one important
group of structures and junctions within one subdivision of the plant kingdom,
evolution has proceeded by means of the type of mutation which genetic study
has shown to be the prevalent type. In connection with this particular phylo-
genetic problem, the familiar difficulty of reconciling ““progressive evolution”
with genetic results does not arise.

It may be added that the mutative changes here shown to have been
important are in harmony with the tendency toward the sterilization of sporo- —
genous tissue which has characterized the evolution of bryophytes, pterido-
phytes, and seed plants.

The succession of steps in the changes from primitive hermaphroditism
must remain for the present-speculative. Obviously male sterility and female
sterility may appear in different plants of a single species, as has happened
in Rubus (4). In Rubus, however, dioecism is not yet reached, for matings
of certain male and certain female plants produce some hermaphrodites and
some “‘neuters” (without functional stamens or pistils). The species may at
present (not considering the neuters) be classed as trioecious. At least two
additional genetic changes would seem to be necessary (11) in order ior
the ultimate goal to be reached. Since mutations are likely to occur inde-
pendently, it is to be expected that in the transition from hermaphroditism
species now dioecious have passed through several intermediate stages.

It is possible to imagine the early steps to have been by way of gyno-
or andromonoecism, trimonoecism, or monoecism. Any of these conditions
could conceivably be reached by the establishment in homozygous condition

AEEEN: PVOLUTION OF SEXUAL CHARACTERS 13

of one or (more probably) more than one mutant gene. How dioecism may
arise from monoecism is illustrated by the success of Jones (8, 9) and of
Emerson (6) in the production of dioecious races of maize through the selec-
tion of appropriate mutations. In each case two mutations were involved;
and in each of the three dioecious races obtained, one pair of chromosomes
differed with respect to a mutant gene which is epistatic to the mutant allele
of the other selected pair.

Another conceivable transition from hermaphroditism to dioecism is by
way of gynodioecism, which has been considerably studied, or of andro-
dioecism, about which nothing is known genetically. If within a hermaphro-
ditic species male sterility becomes a fixed character of one strain, evidently
other strains of the species must usually retain functional pistils if the species
is to persist—that is, a condition of gynodioecism must ensue. An alternative
would be the development (by an additional mutation) of a structurally
female but functionally parthenogenetic species. This seems to have happened
in Hieracium excellens (14) ; but such mutational coincidences must be rare.
A mutation (or mutations) transforming the hermaphrodites of a gyno-
dioecious species into males would lead to dioecism.

The difficulty of explaining the behavior of gynodioecious species by any
simple genic scheme led Wettstein (18) to the assumption of a cytoplasmic
influence—an idea tentatively accepted by Correns (3) and recently empha-
‘sized and generalized by Lewis (10, 11). Apart from the inadequacy of an
explanation based upon one or two mutations, the argument for a cytoplasmic
inhibition in the female upon the functioning of male-tending genes rests
upon the demonstration of such an apparent influence in several typically
hermaphroditic plants, including forms of Linum, Nicotiana, Geranium,
Epilobium, and Streptocarpus. WWith the exception of one case in maize (15),
the known phenomena of this nature are limited to interspecific hybrids. On
the other hand, also, gene mutations leading to male sterility are, as has been
seen, of frequent occurrence. It is entirely possible that, when the variable
behavior of gynodioecious species becomes better known, a (perhaps com-
plicated) Mendelian explanation may be found possible.

Nearly twenty years ago Emerson (5), pointing out that “there are at
least nine pairs of genetic factors which influence the expression of sex in
maize,” suggested that “the genetic situation in maize ... . may perhaps
afford some help toward a solution of sex problems.” The prophecy has
been abundantly confirmed. Today more than 40 genes are known in maize,
borne on at least 9 of the 10 chromosomes, whose presence in the “normal”
or usual condition is directly essential to the sex-expression typical of the
species as it exists at present. There are others, likewise essential in this
regard, whose more conspicuous influence is upon the form, size, or vigor

14 AOR RE Y A

of the plant. There can be no serious doubt that the sex-expression of other
angiosperms, many of whose mutations parallel those observed in maize,
is likewise dependent upon the activity of many genes.

A large proportion of the genes concerned in the sex-expression of con-
temporary species may have come down unchanged or little changed from
primitive angiosperms. Those early species were themselves the outcome
of a long evolutionary history in whose course had been developed a complex
genotype. As has been seen, the passage from hermaphroditism to other
sexual conditions need involve only a comparatively small number of muta-
tions of genes already present. One step—perhaps in general the final step
if maize may serve as an example—involved the establishment in some mem-
bers of heterozygosis with reference to one pair of genes as to which other
members of the species are homozygous. The pair of chromosomes bearing
this allelic pair now plays a part in sex-determination.

Again to judge from maize, the selection of different mutant genes may
in different cases give rise to the same phenotypic result—namely, dioecism.
This example shows, too, how different pairs of chromosomes may in dif-
ferent cases come to function in sex-determination, as the X-Y pair appears
to function in some seventy-odd species of dioecious angiosperms. The rela-
tion of the sex chromosomes to the differentiating genes may vary from
species to species. In Rumex (13), as in Drosphila, the Y chromosome plays
no demonstrable part in sex-differentiation. In Lychnis, on the other hand
(17), its role is a positive one. In Fragaria, as in one of Emerson’s derived
races, the “X-Y” pair characterizes the female of the species; in all other
well-known cases in angiosperms, this pair is the property of the male. While
a partial picture is thus presented of the functioning of a pair of chromosomes
in sex-separation, no satisfactory explanation is yet available for the frequent
visible differentiation between the members of this pair. At the same time,
it is shown in more than forty investigated angiosperms that there is no
necessary correlation between the final genic differentiation which in a
dioecist influences sex and a perceptible difference in chromosome size or
appearance.

The mutations that have determined the transition from hermaphroditism
have not produced in most cases, if in any, an absolute fixity of sexual char-
acter. Instead, whatever the inhibiting tendencies of a particular mutation,
it remains possible, under favoring conditions, for some or all of the old
potentialities to be manifested ; as when bisexual flowers appear on a monoecist
or dioecist. No rigidity of sex-separation seems to have been reached by the
angiosperm sporophyte such as characterizes the angiosperm gametophyte
or the gametophyte of a dioecious bryophyte.

The conception which emerges of the genetic basis for sex can not be
satisfactorily formulated in terms of so many genes for maleness and so

ALLEN: EVOLUTION OF SEXUAL CHARACTERS 15

many for femaleness. The elements which constitute this mechanism at any
particular period in the history of a species influence in various ways and in
varying degrees the development and functioning of stamens and pistils ; they
influence also the numbers and arrangement of these organs, whether in the
same or in separate flowers, as well as the time of appearance of the respective
flowers and their positions on the plant.

DEPARTMENT OF BOTANY
UNIVERSITY OF WISCONSIN
Maprson, WISCONSIN

Literature Cited

Most of the studies referred to in the present paper are cited in the first article listed
below. For this reason only a few references to specific papers have been included.

1. Atten, C. E. The genotypic basis of sex-expression in angiosperms. Bot. Rey. 6: 227-
300. 1940.

2. CorreNns, C. Die Bestimmung und Vererbung des Geschlechtes nach neuen Versuchen
mit hoheren Pflanzen. 1907.

. Bestimmung, Vererbung und Verteilung des Geschlechtes bei den hoheren
Pflanzen. Handbuch der Vererbungswissenschaften (Baur and Pasenann) Int. (G
1928.

4. Crane, M. B., and W. J. C. Lawrence. The genetics of garden plants. 2d ed. 1938.

5. Emerson, R. A. A genetic view of sex expression in the flowering plants. Science
N. S. 59: 176-182. 1924.

6. ————. The present status of maize genetics. Proc. 6th Int. Congr. Genetics 1: 141-152.
1932.

, G. W. Beapte, and A. C. Fraser. A summary of linkage studies in maize.
Cornell Univ. Agr. Exp. Sta. Mem. 180. 1935.

8. Jones, D. F. Dioecious maize. Science N. S. 73: 432. 1931.

. Unisexual maize plants and their bearing on sex differentiation in other plants
and in animals. Genetics 19: 552-567. 1934.

10. Lewis, D. Male sterility in natural populations of hermaphrodite plants. New Phytol.
40: 56-63. 1941. }

. The evolution of sex in flowering plants. Biol. Rev. 17: 46-67. 1942.

12. Nacat, J. Studies on the mutations in Oryza sativa L. I-IV. Jap. Jour. Bot. 3: 25-96.
1926.

13. Ono, T. Chromosomen und Sexualitat von Rumex Acetosa. Sci. Repts. Tohoku Imp.
Univ. 4th Ser. Biol. 10: 41-210. 1935.

14. OstenFeLD, C. H. Castration and hybridisation experiments with some species of
Hieracia. Bot. Tidskr. 27: 225-248. 1906.

15. Ruoapes, M. M. The cytoplasmic inheritance of male sterility in Zea Mays. Jour.
Genet. 27: 71-93. 1933.

16. Suneson, C. A. A male sterile character in barley. Jour. Hered. 31: 213, 214. 1940.

17. Warmke, H. E., and A. F. BLAKESLEE. Sex mechanism in polyploids of Melandrium.
Science N. S. 89: 391, 392. 1939.

18. Werrstetn, F. Uber Fragen der Geschlechtsbestimmung bei Pflanzen. Naturwiss.
12: 761-768. 1924.

19. Werrstern, R. Handbuch der systematischen Botanik. 4th ed. 1935.

20. Yampotsky, C., and Herene YAmporsky. Distribution of sex forms in the phanero-
gamic flora. Biblioth. Genet. 3: 1-62. 1922

VoL. 43 TOUR RAE YO Jury 1943

Leaf-stem Relationships in the Vascular Plants*

RatpH H. WETMORE

It is an arresting fact that in the year of our Lord 1942 there is still no
general agreement on the organization of the Vascular Plants. Perhaps the
nearest one can get to a generalization is the admission that Vascular Plants
ordinarily comprise root systems and shoot systems. Studies of the shoot
systems indicate the usual presence of stems, leaves and reproductive parts.
The interpretation of the relation of leaves to the stem which bears them has
been varied. From time to time there have been those who adhered to the
phyton hypothesis, a hypothesis that made the leaf the important unit of con-
struction of the shoot system, each leaf consisting of the foliar appendage
and its subjacent stem segment or internode. This concept by which the stem
becomes a vertical aggregation of leaf bases was probably advocated first by
Gaudichaud in 1841, and subsequently by Schultz (1843) and by Delpino
(1880, 1883). Little of really scientific contribution could be attributed to
these workers. Their fanciful ideas, however, were given an artificial bolstering
by Celakovsky (1901) when he brought toegether a group of serious arguments
supporting the foliar nature of the stem. However, as Schoute (1931) points
out these same facts upon which Celakovsky’s arguments were based could
equally well be explained otherwise. This early hypothesis did not have a
large or literal following. No more did Chauveaud’s phyllorhiza hypothesis
(1921) a modification of the phyton concept, meet with general acceptance.

Alternative ideas, which held the stem to be an independent organ bearing
foliar appendages, have been prominent and generally much more in favor.
Our textbooks bear witness to this fact.

With these two concepts of the shoot system in mind, I should like to
present the results of certain recent and current developmental studies. Since
the work of Buder (1928) and his students (Schmidt, 1924, etc.) on apical
meristems there has grown a body of knowledge which challenges the form-
alized interpretations of developmental patterns in the apices of root and
shoot attributed to Hanstein (1868). These studies have been continued
especially by the significant works of Schtiepp (1926) and Foster (1935, 1936,
1938, 1939a, 1939b, 1940, 1941a, 1941b). The latter investigator has made
progress on a comparative study oi the diverse types of apical meristems in
different plant groups. The already classical works on Helm (1931, 1932)
and Louis (1935) have extended our knowledge of the whole stem tip
with its developing leaves. More recently numerous workers have contributed

* Read at the 75th Anniversary Celebration of the Torrey Botanical Club at Columbia
University, Monday, June 22, 1942.
i 16

WETMORE: LEAF-STEM RELATIONSHIPS 17

to this field—Priestley (1928, 1929, 1935, 1936) and his associates (Griffiths
and Malins, 1930; Majumdar, 1942; Scott and Priestley, 1925), Esau (1938,
1939, 1940, 1942, 1943), Cross (1937, 1939, 1940, 1941, 1942) to name a few.
These studies bear directly on any interpretation of the leaf-stem relationship.

Louis reported in detail on the development of the stem apices in nine
Angiosperms and one Gymmosperm—Tarus baccata. In his study of Syringa
vulgaris, with opposite leaves, he called attention to the general flat appearance
of the apical meristem upon which the paired leaf primordia are elevated
(Fig. 1). In their early appearance these peg-like protuberances are composed
of cells like those of the meristem itself. At this level a transverse section of
the tip indicates an oval or near rectangular shape, with each end of the
rectangle called a leaf buttress——Majumdar’s leaf foundation (1942)—on
which originates an erect leaf primordium. Such a transverse section Louis
(following Schmidt, 1924) considered as made through a region of maximal
area (Fig. 2). If another transverse section be cut immediately below this pair
of leaf primordia, a circular outline is obtained ; Louis considers such a circular
section as cut through a reg.on of minimal area (Fig. 3). A section through
the bases of the next lower pair of leaves provides another region of maximal
area, with its major axis at right angles to the first. Thus the stem tip is
composed of alternating zones of maximal and minimal area, each buttress
of the former always bearing a leaf primordium. Obviously then a leaf buttress
is topographically a part of the stem with no clear boundary between it and its
elevated leaf primordium.

Studies of successive leaves proceeding downward from the apex give one
a progressive picture of developmental changes in the leaf. As Louis points
out, such studies point to a general increase in vacuolation on the outer or
abaxial side of each young primordium and its buttress. Shortly thereafter in
each leaf an adaxial area appears as equally vacuolated. Thus in Syringa there
is left between the two vacuolated areas, continuous with the apical meristem, a
band of tissue as seen in transverse section ( Figs. 2, 3, 4).

Careful examination shows this band to be heterogeneous, and comprised
of a leaf bundle in procambial stage flanked by residual meristem (Esau, 1943).
As one follows the sections downward it is seen that the first two pairs of leat
buttresses completely surround the stem, at which level the four abaxial ground
meristems, appearing extensively vacuolated, now envelop the stem as the in-
cipient cortex (Fig. 4). Successive sections also show that each adaxial, highly-
vacuolated ground meristem has become continuous with the pith, thereby
forming the so-called leaf gap.

Thus at a level below the second pair of leaves (Fig. 4) the stem of Syringa
consists externally of a protoderm surrounding a potential cortex. A pith is
clearly evident as an early vacuolated, central ground meristem (Fig. 1, 4).

FORREY A

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WETMORE: LEAF-STEM RELATIONSHIPS 19

Between these cortical and pith ground meristems exists a ring of small-celled
tissue which Louis has designated prodesmogen. An examination of this ring,
however, shows it to be heterogeneous in nature, not homogeneous as Louis
supposed. Confronting each leaf buttress is a small-celled arc of differentially
staining tissue—the procambium of the leaf trace—which is continuous with
that of the developing leaf. On either side of this leaf trace bundle, in the ring,
is a narrow zone of residual meristem,—the primary ray. Immediately above
each outward-bending leaf trace is the highly vacuolated local break in the
ring, referred to above as the leaf gap (Fig. 1, right). Louis’ investigation of
Syringa does not include the development of procambium. However subse-
quent study by the writer indicates the continuity of this procambium with
differentiating primary vascular tissue below, its development being con-
tinuously acropetal.

Allowing for variations according with the phyllotactic pattern, size of
leaf and number of leaf traces per leaf in the large number of Angiosperms
now investigated,* it would seem fair to state that in this large group of
plants vascular and cortical patterns are generally correlated with the forma-
tion of leaves at the apex. The pith by contrast seems to belong to the axis.
It was on the strength of such studies in his own laboratories that Priestley
and his associates propounded the idea of “the unit of shoot growth’ for
Angiosperms, a modified phyton and a unit closely resembling the “Spross-
glieder’”’ of Celakovsky. Each such unit consists of a leaf and a subtending
longitudinal sector of the stem—not a whole segment as earlier phytonists had
considered it. There are many interesting points in this hypothesis as Priestley
has developed it. Time does not permit their consideration. It is true that in
the Angiosperms investigated by Priestley the facts could be so interpreted.
It is perhaps equally pertinent to question whether the generalization which
he makes will hold for all cases in the Angiosperms. In this connection I
should like to call your attention to Hippuris vulgaris, which Louis has also

* Studies by Foster (1938, 1939b, 1940, 1941la, 1941b) and his student Gifford (1943),

Crafts (1940), Cross (1939, 1940, 1941), etc., would indicate that this statement is perti-
* nent also for Ginkgo, the Cycads, the Conifers and Ephedra.

Explanation of figures 1-5

Fics. 14. Syringa vulgaris. Fig. 1. Longitudinal section of stem tip (X90).
Fig. 2. Transverse section of stem tip showing region of maximal area through buttresses
of first pair of leaves (X130). Fig. 3. Transverse section slightly lower through
buttresses of first pair of leaves near region of minimal area; leaf gap almost confluent
with pith (X130). Fig. 4. Transverse section below attachment of first two pairs
of leaves to show the ring composed of procambium and primary rays or interfascicular
residual meristem (130). Fig. 5. Longitudinal section of stem tip of Hippuris
wulgaris (X130). (Figs. 1-5 after Louis.)

i ORRIRGE, YAN

WETMORE: LEAF-STEM RELATIONSHIPS 21

illustrated. As is well known, this plant possesses a protostele in the stem.
The apical meristem is not flat as in most Angiosperms; the leaves are
borne laterally (Fig. 5). There is a central procambial column—i.e., no pith—
which rises above the last leaf primordium. Cortical ground meristem also
exists above the highest leaf primordium. In other words, the stem now shows
a potential epidermis, a potential cortex, and conducting tissues without the
“influence” of leaf primordia.

It may be argued that this aquatic plant is modified in relation to its envi-
ronment. Possibly so; nevertheless, it seems significant that the plant should
be developing at all, if the axis of Angiosperms is a system of phytons or
growth units only. On such slight evidence one can do no more than suggest the
possibility that Angiosperms have a shoot system, potentially both cauline and
foliar, in which ordinarily the leaves possess a dominant and the stems a
minor “influence” on development, but in which on occasion the stem may
hold the major role and the leaves a minor though necessary one. In this
connection one might refer to certain seeming cauline bundles—certainly not
associated with leaves—which Boke (1941) reports in the cactus Trichocereus
Spachianus.

It is instructive to examine other groups of vascular plants for develop-
mental patterns. Basing one’s judgments on the Angiosperms alone may
well produce a limited outlook. May I call your attention to the genus Lyco-
podium in which our laboratories have been interested for some time. The
conclusions are based on a careful investigation of nine species. Those repre-
sentatives which we have studied from the Urostachys segregation of Lyco-
podium, L. lucidulum Michx. and L. Selago L. have a flat-topped apical
meristem with erect foliar primordia (Fig. 6). The protostelic vascular cylinder
is forecast in a recognizable column of procambial tissue which rises higher
toward the apex than the place of origin of the youngest leaf primordium.
Young leaf primordia already show procambial strands related to them, never
discontinuous with the central column. The cortical ground meristem is
belated in appearance. There is never a pith nor is there any adaxial vacuolated
ground meristem with its associated leaf gap. By studying successive trans-
verse sections below the apex it is seen that the cauline vascular tissues—the
metaxylem and metaphloem—are outlined or blocked out, within 100, of
the apex in L. lucidulum, whereas the first sign of the differentiation of pro-
toxylem occurred only about 300p from the apex. However, though blocked

Explanation of figures 6 and 7

Longitudinal sections of serial stem tips of Lycopodium to show proximity of pro-
cambial column to the stem apex. Fig. 6. L. Selago; stem apex flat-topped (X260).
Fig. 7. L. sabinaefolium; stem apex conical (X 300).

22 OUR BVA

out, no differentiation of this cauline tissue into metaxylem and metaphloem is
seen to occur until after differentiation in the leaf traces themselves has
become well established, and then only centripetally from the protoxylem and
protophloem (Fig. 8). It is interesting to recall that such a blocking out
of the metaxylem and metaphloem pattern, before differentiation occurs, is
reported commonly in Angiospermous roots. (Esau, 1940; Williams, 1940.)

Fig. 8. Tranverse section of aerial stem of L. sabinaefoliwm, 2710" trom apex,
showing pattern of the radially organized stele already blocked out, but differentiation
only present in the peripheral strands of protoxylem and protophloem (X 260).

In the remaining part of the genus Lycopodium, as represented by the
seven species studied,* the apex is not flat, but conical, with laterally borne
leaves (Fig. 7). Here, however, the developmental story is generally the
same (Fig. 8). So is it true for the numerous species of Selaginella now in
process of investigation ; the detailed study is not yet complete. ‘Before leaving

* This study includes Lycopodium inundatum L., L. cernuwm L., L. annotinum L.,

L. clavatum L., L. obscurum L., L. sabinaefolium Willd., L. complanatwm L., and its
variety flabelliforme Fernald.

WETMORE: LEAF-STEM RELATIONSHIPS 23

Lycopodium, I should like to mention that all the underground rhizomes of
L. obscurum so far examined have only membranous scale-like leaves with no
traces. It is significant that this rhizome develops vascular tissues and cortex,
however, not dissimilar to those normally present in a leaf-bearing rhizome.

In substance, these living Lycopsida seem to have a shoot system composed
of a cauline part with foliar primordia borne initially in either erect or lateral
position. Here there seems to be a peripheral set of bundles, which form a
primary network and which are connected with the leaves, bundles originating
from procambial strands which are truly acropetal and continuous in their
origin. In addition, ordinarily differentiating only after leaf connections are
established, though blocked out earlier, is the whole central portion of the
central cylinder which is cauline in nature and which is never directly connected
with the leaves.

What of the Horsetails, Ferns, and Gymnosperms? In Equisetum, the
story is far from complete. A study of native species of this genus now in
progress in our laboratories gives evidence of a continuous acropetally develop-
ing procambium to the leaves and branches. It is not yet clear, however, from
this work, nor that of Barratt (1920), Queva (1907) or Vidal (1912), just
how the nodal ring is developed. Certainly it would be difficult to think of
this ring as entirely foliar in nature.

In the Ferns, the study is fragmentary. The work of Gillette (1937)
on Psaronius and of Schoute (1926) on living Marattiaceae suggest the
complicated stele to be of foliar origin. In the three native species of Os-
mundaceae, as yet unreported studies from our laboratories show no sign
of cauline bundles, though Kaplan reports such. In his summary on the Ferns
in Verdoorn’s MANUAL oF PTERIDOLOGY, Schoute (1938) states (p. 84): “In
the Ferns the original Pteridophyte stele with its external sheaths, its phloem
and its central solid xylem has been reduced into a mere topographical tissue
column, acting as a egy piem for leaf-traces, but without any tissue dif-
ferentiation of its own.’

Before leaving the Ferns, I must peter to certain scale-bearing stolons of
species of Nephrolepis, studied by Lachmann (1885, 1889) and others.
Originally described as roots, they proved to be stems with a cortex and a
protostelic central cylinder resembling much more that of sporeling Ferns
or mature axes of Gleichnia, Lygodium or Hymenophyllum which remain
permanently protostelic.

In the Gymnosperms, the classical account of Koch (1891), followed by
those of Barthelmess (1935), Cross (1939, 1940, 1941, 1942), Foster (1938,
1939b, 1940, 1941a, 1941b), Gifford (1943), Korody (1937), and Louis
(1935), indicates diverse patterns of apical meristem in the different gymno-
spermous assemblages. The work is too incomplete to give any summary

24 TORRONE DY A

statement. Barthelmess (1935), in his study of various Conifers, considers
the primary vascular tissue composed of leaf traces only, variously united
into sympodia. However, his interpretation of procambium developing basi-
petally is not in agreement with Cross’ findings (1942) in Cunninghamuia
lanceolata, Crafts (1940) in Sequoia, and our as-yet unpublished findings in
Pinus Strobus and Ginkgo biloba. In general, Barthelmess points out the
similarity of the coniferous apical region to that described for the Angio-
sperms by Helm (1931). One observation of Barthelmess’ should be referred
to, that of a shoot of Pseuwdotsuga which in the course of its development failed
to produce the normal needle-like leaves and instead gave only membranous,
scale-like structures with no leaf traces. Yet this shoot when examined gives
a normal structural picture for a shoot of Pseudotsuga except that the vas-
cular cylinder is unbroken by the usual interfascicular parenchyma or primary
rays.

A summary survey of the literature and current research pertaining to
the organization of primary shoots of diverse groups of the vascular plants
certainly leaves the writer with no final dictum on the nature of the shoot.
There is cumulating evidence, however, that in Lycopodium and Selaginella
the vascular cylinder is mostly of cauline, to a lesser degree of foliar, origin.
Even here, though blocked out somewhat earlier in development, the differen-
tiation of metaxylem and metaphloem ordinarily does not seem to occur until
the leaf traces are themselves in a process of differentiation.

The evidence for Equisetum, Ferns, and Gymnosperms is still too incom-
plete to permit of generalizations. The rhizomes of Lycopodium obscurum,
the leafless stolons of Nephrolepis, the unusual shoot of Pseudotsuga, and
other leafless cauline axes considered by Troll (1937, p. 287-304), give
indication that stems in these groups may develop epidermis, cortex and vas-
cular tissues even though no leaves be present. Certainly the early appearance
of pith in the majority of species of Horsetails, Ferns, and Gymnosperms
would suggest for it a cauline origin, for this pith is often found higher in the
axis than the most apical, foliar, procambial connection. As Schoute ( Verdoorn,
1938) points out in his summary for the Ferns, there can be no question as to
the “influence” of megaphyllous leaves on the differentiation of vascular tissues.

In the Angiosperms, developmental studies generally give indication of
the importance of foliar structures on the entire developmental sequence of
events in the axis. There is little evidence to suggest a separate role for the
axis in the development of vascular tissues, except possibly in certain aquatics
such as Hippuris, possibly in the interesting case of the cactus Trichocereus,
and a few other instances. However, the fact that roots develop vascular and
cortical tissues without foliar appendages must not be forgotten.

Is it possible to consider the shoot system as an entity within which a

WETMORE: LEAF-STEM RELATIONSHIPS 25

division of labor has occurred, the leaf being set off physiologically from the
axis bearing it even though it originated as a product of the same meristematic
activity which adds to the stem tip? There is increasing evidence that in
many cases in diverse groups of Vascular Plants each foliar primordium so
produced is provided with procambium continuous from below at all times
(Esau, 1943; Wetmore and Smith, 1942, etc.). Whatever the later orderly
differentiation of primary xylem and primary phloem may be, that continuity
seems of significant import. The boundary then between leaf and axis is
indefinite with leaf buttresses present as those parts of the axis from which
leaf primordia are elevated. It must be pointed out, however, that the
“influence” of the leaf is of different degrees in different groups of plants.
Certainly in roots, in rhizomes of Lycopodium obscurum, in leafless axes of
Nephrolepis, and in the leafless shoot of Pseudotsuga, cortex is produced
as well as a vascular cylinder. In the cauline structures, the cortex is ordinarily
retarded in its development. In Lycopodium and Selaginella, microphyllous
plants, cortex is ordinarily slow in developing. In the Conifers, one finds
needle-like or scale-like small leaves and a slowly developing cortex. In the
Angiosperms and Ferns with their characteristic large leaves, cortical and
vascular differentiation is early, yet small-leaved types such as Linum show
the usual delay (Esau, 1942). As Kaplan (1937) has suggested, cortex
appears soonor or later but leaves seem to accelerate the process of cortical
vacuolation.

If I, this early, should venture to epitomize the leaf-stem situation, it would
be something as follows: The early, psilopsid land plants, still leafless, were
protostelic. With the advent of leaves, microphyllous or megaphyllous, various
changes have occurred in stem organization. Microphyllous plants possess
in their primary axes a small amount of “foliar” trace vascular tissue, periph-
erally connected to the cauline, vascular cylinder. In megaphyllous groups,
the foliar vascular system and its stem connections become more significant
and the potential cauline portions, failing in varying degrees to differentiate,
appear as pith. The shoot system is the sum total of foliar and cauline expres-
sion. From the practical point of view, the shoot is still composed of leaves
borne on a stem system. From a developmental point of view, where one of
necessity is faced with factors underlying development, an understanding of
the varied developmental patterns of shoot expression in the vascular plants
seems significant. How else can one approach experimentation to determine
the underlying physiological and biochemical background than with a knowl-
edge of the structural and developmental variables ?

THE BroLocicaL LABORATORIES

Harvarp UNIVERSITY.
CAMBRIDGE, MASSACHUSETTS

26 GO IRSRGE CY IAs

Literature Cited

Barratt, Kate. 1920. A contribution to our knowledge of the vascular system of the
genus Equisetum. Ann. Bot. 34: 201-235.

BartHELMEss, A. 1935. Ueber den Zusammenhang zwischen Blattstellung und Stelenbau
unter besonderer Bertichsichtigung der Coniferen. Bot. Arch. 37: 207-260.

Boxe, N. H. 1941. Zonation in the shoot apices of Trichocereus spachianus and Opuntia
cylindrica. Amer. Jour. Bot. 28: 656-664.

Buper, J. 1928. Der Bau des phanerogamen Sprossvegetationspunktes und seine Bedeu-
tung ftir die Chimarentheorie. Ber. Deut. Bot. Ges. 46: (20)-(21). ‘

CEeLAKovskY, L. J. 1901. Die Gliederung der Kaulome. Bot. Ztg. 59: 79-114.

CHAUVEAUD, G. 1921. La constitution des plantes vasculaires révélée par leur ontogénie.
Payot et Cie, Paris.

Crarts, A. S. 1940. Vascular differentiation in the shoot apex of Sequoia. Amer. Jour.
Bots 2h AOD:

Cross, G. L. 1937. The morphology of the bud and the development of the leaves of
Viburnum rufidulum. Amer. Jour. Bot. 24: 266-276.

. 1939. The structure and development of the apical meristem in the shoots of

Taxodium distichum. Bull. Torrey Bot. Club 66: 431-452.

. 1940. Development of the foliage leaves of Tarodium distichum. Amer. Jour. Bot.

27: 471482.

. 1941. Some histogenetic features of the shoot of Cryptomeria japonica. Amer. Jour.

Bot. 28: 573-582.

. 1942. Structure of the apical meristem and development of the foliage leaves of
Cunninghamia lanceolata. Amer. Jour. Bot. 29: 288-301.

Cross, G. L. and T. J. Jounson. 1941. Structural features of the shoot apices of diploid
and colchicine-induced tetraploid strains of Vinca rosea L. Bull. Torrey Bot. Club 68:
618-635. ;

Dexpino, F. 1880. Causa meccanica della fillotassi quincunciale. Nota preliminare. Genova.

. 1883. Teoria generale della fillotassi. Atti della R. Universita di Genova 4.

Esau, K. 1938. Ontogeny and structure of the phloem of tobacco. Hilgardia 11: 343-424.

. 1939. Development and structure of phloem tissue. Bot. Rev. 5: 373-432.

. 1940. Developmental anatomy of the fleshy storage organ of Daucus carota. Hil-

gardia 13: 175-226. :

. 1942. Vascular differentiation in the vegetative shoot of Linum. I. The procam-

bium. Amer. Jour. Bot. 29: 738-747. é

. 1943. Origin and development of primary vascular tissues in seed plants. Bot.
Rev. 9: 125-206.

Foster, A. S. 1935. A histogenetic study of foliar determination in Carya Buckleyi var.
arkansana. Amer. Jour. Bot. 22: 88-147.

. 1936. Leaf differentiation in angiosperms. Bot. Rev. 2: 349-372.

. 1938. Structure and growth of the shoot apex in Ginkgo biloba. Bull. neeaen Bot.

Club 65: 531-556.

. 1939a. Problems of structure, growth, and evolution in the shoot apex of seed

plants. Bot. Rev. 5: 454-470.

. 1939b. Structure and growth of the shoot apex of Cycas revoluta. Amer. Jour.

Bot. 26: 372-385.

. 1940. Further studies on zonal structure and growth of the shoot apex of Cycas

revoluta Thunb. Amer. Jour. Bot. 27: 487-501.

WETMORE: LEAF-STEM RELATIONSHIPS 27

. 1941a. Comparative studies on the structure of the shoot apex in seed plants.

_ Bull. Torrey Bot. Club. 68: 339-350.

. 1941b. Zonal structure of the shoot apex of Dioon edule Lindl. Amer. Jour. Bot.

28: 557-564.

. 1942. Practical Plant Anatomy. D. Van Ostrand Co., New York.

GaupicHaAup, Cu. 1841. Recherches générales sur l’organographie la physiologie et l’or-
ganogénie des vegetaux. Paris. .

GrrForD, Ernest M. 1943. The structure and development of the shoot apex of Ephedra
altissuma Desf. Bull. Torrey Bot. Club. 70: 15-25.

GitteTTE, N. J. 1937. Morphology of some American species of Psaronius. Bot. Gaz. 99:
80-102.

GrirFitHs, A. M. and M. E. Matins. 1930. The unit of shoot growth in dicotyledons.
Leeds Phil. and Lit. Soc., Proc., Sci. Sec. 2: 125-139.

Hanstein, J. 1868. Die Scheitelzellgruppe im Vegetationspunkt der Phanerogamen.
Festschr. Niederrhein. Ges. Natur- u. Heilkunde. 109-143.

Heim, J. 1931. Untersuchungen iiber die Differenzierung der Sprosscheitelmeristeme von
Dikotylen unter besonderer Berticksichtung des Procambiums. Planta 15: 105-191.

. 1932. Uber die Beeinflussung der Sprossgewebe-Differenzierung durch Entfernen
Junger Blattanlagen. Planta. 16: 607-621.

Kapran, R. 1937. Uber die Bildung der Stele aus dem Urmeristem von Pteridophyten und
Spermatophyten. Planta. 27: 224-268.

Kocu, L. 1891. Uber Bau. und Wachsthum der Sprossspitze der Phanerogamen. I. Die
Gymnospermen. Jahrb. f. Wiss. Bot. 22: 491-680.

Koropy, ExisaBetH. 1937. Studien am Spross-Vegetationspunkt von Abies concolor,
Picea excelsa und Pinus montana. Beitr. Biol. Pflanz. 25: 23-59.

LACHMANN, P. 1885. Recherches sur la morphologie et l’anatomie der Fougéres. Compt.
Rend. Acad. Sci. 101: 603-606.

-. 1889. Contributions a l’histoire naturelle de la racine des Fougéres. Thesis. Lyon.

Louts, JEAN. 1935. L’ontogénése du systeme conducteur dans la pousse feuillée des Dico-
tylées et des Gymnospermes La Cellule. 44: 87-172.

Mayumpar, G. P. 1942. The organization of the shoot in Heracleum in the light of develop-
ment. Ann. Bot. n.s. 6: 49-81.

PriestLey, J. H. 1928. The meristematic tissues of the plant. Biol. Rev. 3: 1-20.

. 1929. Cell growth and cell division in the shoot of the flowering plant. New Phytol.
28: 54-81.
PriestL_ey, J. H., L. I. Scorr, and E. C. Gitte. 1935. The development of the shoot in
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Priestley, J. H., and L. I. Scorr. 1936. The vascular anatomy of Helianthus annuus L.
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ScHumipt, A. 1924. Histologische Studien an phanerogamen Vegetationspunkten. Bot. Arch.
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Scutepp, O. 1926. Meristeme. In K. Linsbauer’s Handbuch der Pflanzenanatomie. Bd. 4.

Scnoute, J. C. 1926. On the foliar origin of the stelar structure of the Marattiaceae. Rec.
Trav. Bot. Néerl. 23: 269-304.

. 1931. On phytonism. Rec. Trav. Bot. Néerl. 28: 82-96.

Scuuttz, C. H. 1843. Die Anaphytose oder Verjiingung des Pflanzen. Ein Schlussel
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-sichten auf die Cultur der Pflanzen. Berlin.

28 GUOTRIRE ES YA

Scott, L. I., and J. H. Priestrey. 1925. Leaf and stem anatomy of Tradescantia fluimi- -
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Trott, Wm. 1937. Vergleichende Morphologie der hoheren Pflanzen. 1 Band. 1 Teil.
Gebrtider Borntraeger, Berlin.

VeRpDOoRN, Fr. (editor). 1938. Manual of Pteridology. Martinus Nijhoff, The Hague.

VipaL, L. 1912. La croissance terminale de la tige et la formation des bourgeons chez
VEquisetum palustre. Ann. Sci. Nat., Bot., 9¢ sér., 15: 1-38.

Wetmore, R. H., and Apa C. Smiru. 1942. Development of shoot system in Lycopodium.
Amer. Jour. Bot. 29: 19s.

WiiiaMs, Bert C. 1940. Differentiation of vascular tissue in root tips. Amer. Jour. Bot.

27: 10s.

Vor. 43 DO MRIReaD YA Jury 1943

Cell Division as a Problem of Pattern in Plant Development*

Epmunp W. SINNOTT

The plane in which a cell divides and the position of the new wall laid
down between the two daughter cells involve important problems, not alone
as to the behavior of individual cells, but also as to the development of multi-
cellular plant structures, since the planes of division in a mass of growing
tissue must evidently be related to the direction in which growth occurs, and
thus to the form of the organs produced.

Various hypotheses have been suggested as to factors which determine
the position of the new wall in a dividing cell. Hofmeister showed that such
walls are usually formed at right angles to the longer dimensions of the cell.
Sachs observed that a new wall tends to meet the old one at right angles. The
direction of mechanical pressure, light, electrical currents, and gradients
of various chemical substances have been shown to affect the orientation of
the division wall. Errera and Berthold, later supported by D’Arcy Thompson
and others, maintained that since cell walls in embryonic tissues are thin
and semi-liquid, their position is governed by molecular forces and will be
such that minimum surface and maximum stability result, so that no more
than three walls meet at one point. All these “rules” can be abundantly illus-
trated from plant material, but every histologist has seen exceptions to them.
Some of these have recently been discussed by the writer and Dr. Bloch.
The problem is evidently too complex to be explained by any one hypothesis.
It has too often been approached simply as a question relating to the activity
of single cells rather than of these cells as members of an organized multi-
cellular system. The present paper reports a study of cell division as it occurs
in a simple plant structure, in an attempt to determine what relation there may
be between the manner in which a cell divides and the position which it
occupies in such an organized entity.

The shoot axis of Equisetum provides particularly good material for
such a study. Its growth is centered in a single apical cell and the lineages
of cells arising from this are relatively easy to follow. The structure of the
axis is without serious complication and the leaves are small, simple and in
whorls. A number of previous studies have been made on various species of
this genus and for many of them the development of the shoot apex is well
known. It seems worth while, however, to examine the facts for a single

* Presented at the 75th Anniversary Celebration of the Torrey Botanical Club at Colum-
bia University, Monday, June 22, 1942.

The writer wishes to express sincere appreciatian to his colleague Dr. Robert Bloch,
who carried out the technical part of the study here reported and prepared the illustration

Oo
j=)

ROTOR BY A

species in some detail from the particular point of view of the problem of cell
division.

Equisetum hyemale, one of the larger of our native species, was chosen
ior study. This has a rather massive meristematic region as compared with
some of the more delicate types. Transverse and longitudinal sections, both
median and tangential, were cut through the stem tips which had been coi-
lected at various times from March until June, the period when meristematic
activity and growth are best studied. Cell divisions occur not only at the apex
of the meristem near the apical cell, but for some distance back along the
axis during the differentiation of various tissues.

The manner in which division takes place was found to differ markedly
depending on the location and character of the cells concerned. Some of the
types are as follows:

The large apical cell cuts off a daughter cell from each of its three inner
faces, successively. The new wall is approximately parallel to the old so
that the two daughter cells are dissimilar in shape and usually in size (Fig. 1).

The lower cell elongatés, as seen in longitudinal section, and divides
periclinally. The inner of its daughter cells contributes, by rather irregular
divisions, to a mass of tissue just below the apical cell. The outer one divides
anticlinally, and thus parallel to its long dimension. This portion of the meri-
stem thus consists of a surface layer of elongate cells and an inner mass of
irregularly-shaped ones (Fig. 1).

Most subsequent divisions in the outer layer are anticlinal, thus violating
Hofmeister’s rule, with the new division wall straight and parallel with the
sides of the mother cell. Occasionally, however, usually at the point where a
new leaf primordium will develop, the inner edge of the phragmoplast begins
a straight course but before it reaches the end of the mother cell it swerves
to one side, usually in the basal direction, until it meets the anticlinal wall of
the old cell (Fig. 2). The smaller of the two daughter cells thus produced,
somewhat V-shaped in section, will form the apical cell of one of the leaf
primordia which begin to appear a little further back. Such a wall as here
described is neither across the shorter dimension of the cell nor does it con-
form to a least-surface configuration, although in the inner part of its course
it becomes curved. / |

Shortly below this level there may be seen in longitudinal section a series
of cell divisions across the axis. This marks the beginning of one of the
diaphragms which is such a conspicuous feature of the stem anatomy of
Equisetum. These divisions are always approximately at right angles to the
axis regardless of the particular shape of the cells in which they occur and,
therefore, occupy positions in these cells which violate many of the “rules.”

SINNOTT: CELL DIVISION AS A PROBLEM OF PATTERN 31

Such divisions continue until a considerable amount of diaphragm tissue is
formed (Fig. 3).

The point where each incipient diaphragm meets the outer surface of the
meristem marks the base of a whorl of leaf primordia. While these are still

Fic. 1. Section through the extreme tip of the growing shoot of Equisetum hyemale,
showing apical cell, surface layer of elongate cells, central mass of irregular cells, and
young leaf primordia.

Fic. 2. An oblique division in one of the surface cells. The smaller one will produce
an apical cell of a leaf primordium.

very small there begins to be differentiated within the base of each a series of
provascular strands from which the circle of vascular bundles will later
develop (Fig. 3). Each of these strands arises by a series of longitudinal
divisions (thus at right angles to those in the diaphragm) in a cell row near
the surface of the meristem. These divisions, like those in provascular tissue
generally, run parallel to the long axis of the cell.

In the subepidermal layer of the meristem, along the future ridge of the
axis, where the photosynthetic tissue will later develop, the method of division
1s still different. Here the anticlinal longitudinal walls in a given ceil are in

32 RYOARARSE YS

Fig. 3. Photograph through a lower region of the meristem showing parts of several
diaphragms, two leaf bases and the beginnings of provascular tissue in stem and leaf.

Fic. 4. Three stages in the development of a row of aerenchymatous tissue. Most of
the division walls are opposite those in vertically adjacent cells.

SINNOTT: CELL DIVISION AS A PROBLEM OF PATTERN 33
most cases exactly opposite similar walls in the cells above and below, so that
in such a group of cells, as seen from its outer surface, four walls usually
meet at a point instead of three as in most tissues. The point where these cells
come together is evidently subject to a good deal of strain as the cells expand
and intercellular spaces thus develop here very early (Fig. 4). These become
much enlarged in the mature tissue. The relation between such a type of cell
division and the development of aerenchyma has previously been pointed out.
by the writer.

The surface cells which are to give rise to stomata undergo a remarkable
series of divisions. In a vertical row of cells, every second one is a stomatal
mother cell. The first division in it is longitudinal and usually unequal, with
the new wall convex toward the smaller daughter cell. The next, in the larger
cell, is convex in the opposite direction, so that a lens-shaped cell has now
been cut out with a larger cell on either side. The lens-shaped cell then divides
into two guard cells.

This diversity in the type and direction of cell division in developing plant
tissue is of course not confined to Equisetum but is a familiar feature of the
process of differentiation in all multicellular plants. The important fact which
it emphasizes is that no single method of division is universal, and that every
“rule” is frequently broken. Evidently many factors may be concerned with
determining the plane of cell division. What a given cell will do depends not
upon some general principle of division, common to all cells, but upon the
conditions which exist at that particular place and time. Every cell is a part
of a general developmental pattern, and not only in the way it divides but in
every other aspect of its behavior it seems to be governed by its particular
place in that pattern. Driesch nearly half a century ago summed this up in
his famous aphorism that “the fate of a cell is a function of its position,” and
Vochting many years before said the same thing in almost the same words.

This general fact of development, so well illustrated by the controlled
diversity of mode of cell division in the meristematic tissues of plants, should
be recognized by all students of morphogenesis. In a search for the mechanisms
which operate in the remarkable processes of organic development, we tend
to oversimplify the problem and to postulate factors which have a specific
method of operation. Thus the role of auxin, of light, or of a given gene is
often assumed to be a definite and invariable one, whereas its effect actually
is dependent in very great measure on the internal and external environment
in which it operates. In an eagerness to find specific organ-forming substances
and stimuli we have too often neglected the complex reaction system, the
developmental pattern in which these must work. Knowledge about specific
factors is very useful and is rapidly accumulating, but far more important

3+ EGR RE Vx

would be an understanding of the complex organic system in which they work
and which determines their effect. About this we still know very little.

The problem may perhaps be stated somewhat more vividly by comparing
the operation of a developing organic mechanism with that of others more
familiar to us. A nickel inserted into a slot, for example, will activate a
turnstile or a juke box or a coin telephone. There is but little specificity in the
“stimulus” but a great deal in the mechanism which it activates. If one knew
everything about nickels and how they differ from other coins he still would
fail to understand how a nickel could produce these results, for an answer
to this question lies in the character of the reacting mechanism. In somewhat
the same way, auxin produces one effect in one part of the developing organism
or under one set of conditions, and quite another elsewhere ; and the principle
of minimal surfaces may determine the position of new cell walls at one region
oi the meristem but may be overruled by other factors in another.

The developing organism is a patterned whole, the parts and activities of
which derive their particular character from their relation to this whole,
and should be studied in this relation and not only as independent structures
or processes. An understanding of this organized pattern and the way in which
it controls development and differentiation is the chief task of the science of
morphogenesis.

Ossorn BotanicaL LABORATORY

Yate UNIVERSITY
New Haven, CoNNECTICUT

VoL. 43 TORR EY A JuLy 1943

Contributions of the Torrey Botanical Club to the Development of
Taxonomy*

H. A. GLEASON

Travel back in your mind to 1867. Andrew Johnson occupies the White
House at Washington. Carpet-baggers are rampant in the South. Boss Tweed
has his thumb on the city of New York. Millions of buffalo graze the plains
of Kansas. The first transcontinental railway has not been completed.

And what of science in this country? Botany is still regarded as a proper
subject of study in a ladies’ seminary. Of plant physiology there is none,
although a young Maine physician, George Goodale, may be musing on the
subject. Of plant pathology there is none, although a country school teacher,
Charles Peck, a storekeeper, Benjamin Everhart, and a farmer, Job Ellis, are
actively collecting fungi, and a young medical student, William Farlow, is
beginning an interest in the subject. Of genetics there is none, although there
is a great deal of talk about a recent book called the Origin of Species. No
ordinary college student has yet peeked through a microscope as a part of
his regular classwork, but a sophomore at Michigan Agricultural College,
Charles Bessey, is wishing that he could and a few years later gave the oppor-
tunity to his own students.

In taxonomy conditions are very different. Three distinguished botanists
stand out above all the rest for their taxonomic research, Gray of Cambridge,
Torrey of New York, and Engelmann of St. Louis, although measured by
influence on the teaching and study of botany and consequently by their inspira-
tion of another generation, Torrey and Gray must divide their honors with
another New York man, Alphonso Wood. The plants of the eastern states are
already thoroughly known and no one gives much attention to this region.
In the south Chapman is still discovering undescribed species, and in the
unsettled and largely uncivilized west several adventurous botanists are sending
east large quantities of new material to Gray, Torrey and Engelmann.

In New York, Professor Torrey was the only research botanist, but there
were several young folks who were interested in plants, who liked to tramp
over the hills, along the beaches, or through the pine barrens. These young
folks met with Professor Torrey, exhibited their botanical treasures, recounted
the adventures of their trips, and rejoiced together over the collection of
some uncommon species. Torrey did not encourage them to work for a doctor’s
degree or require them to register for formal courses in botany. He did not
advise them, to explore the jungles of the tropics, where new species could

* Read at the 75th Anniversary Celebration of the Torrey Botanical Club at The New
York Botanical Garden, Tuesday, June 23, 1942.
35

36 TORREYA

be found, or to monograph the genus Carex. Wise in proportion to his years,
he knew that good taxonomists can develop but can not be forced, and he
probably felt and hoped that from such a group there might arise from time
to time a few taxonomists who, through their deep interest, their keen
observation, and their taxonomic curiosity, would really contribute to the
advancement of science. He, therefore, neither overwhelmed them with his
own knowledge nor belittled their own amateur work, but listened patiently
to the accounts of their adventure, praised them for their discoveries, and by
his geniality and interest encouraged them to further study. These were the
men who organized themselves into the Torrey Botanical Club in 1867.

After the death of Torrey, the Club was left to stand or fall on its own
merits. During the seventies it was held together partly by the common inter-
est of its members, which could be expressed in meetings and field excursions,
and partly by the responsibility of publishing the Torrey Bulletin.

As the first contribution which the Torrey Club has made to taxonomy,
we naturally think of its publications. For many years the largest item in the
budget of the Club has been for the production of the BULLETIN, the MEmorrs,
and Torreya. And as the Club has been generous, so have taxonomists, not
only the members of the Club but non-members as well, been fortunate in
finding in it a dignified and reputable means of presenting their results
to the world.

Those who have had occasion to look through the early volumes of the
BULLETIN know that the membership of the Club was originally composed
almost entirely of amateur taxonomists, of young men interested in the local
flora, and that Dr. Torrey was the only professional taxonomist in the group.
From the pens of these young men came a series of short notes, almost all
taxonomic or floristic in nature and most of them very amateurish. Some of
them soon graduated into actual research work; among them T. F. Allen and
C. F. Austin, who began during the seventies to publish critical discussions
and descriptions of new species of Chareae and Hepaticae.

The BULLETIN soon began to attract the attention of other American
botanists, and during the seventies and early eighties its pages contain con-
tributions from such well-known men as F. L. Collins, A. H. Curtis, J. B. Ellis,
George Engelmann, Asa Gray, Charles H. Peck, John Donnell Smith, William
Trelease, L. M. Underwood, and Francis Wolle. As its circulation grew,
so did the length and importance of its articles. Little by little the local
observations disappeared and were replaced by sober research, until during
the eighties and nineties it had become without doubt the leading American
outlet for the publication of taxonomic research. To supplement the BULLETIN
and to provide for longer articles, the Memoirs were established in 1889 and
have given the bulk of their pages also to taxonomy. TorRrEyA was established

GLEASON: CONTRIBUTIONS OF THE CLUB TO TAXONOMY 37

in 1901, primarily for a revival of opportunity for the discussion of local
botany, but it also has given a fraction of its space to taxonomic research.

As a matter of statistics, it may be recorded that to the end of 1941, the
Club has published a total of 22,098 pages of printed matter devoted to pure
taxonomy or to cognate subjects primarily of interest to taxonomists. I feel
certain that this impressive total is not approached by any other American
magazine during the same three-quarters of a century.

In the preparation of this paper, I have leafed through the publications
of the Club and have compiled two graphs showing the amount of taxonomic
publication year by year, and the proportion, expressed in percentage, of the
total publication which has been devoted to taxonomy. In doing so I have
often had to make hasty judgments as to the taxonomic or non-taxonomic
classification of an article, and I have also tried to take into account the general
nature of the membership of the Club and of its audience at the different
periods in its history. Consequently I have included in taxonomy many short
articles from the early volumes which, if printed today, would be regarded
merely as interesting notes of no special botanical value. The resulting graphs,
to revive an ancient New York simile, were as crooked as Pearl Street and
their general trend was badly obscured by the huge annual fluctuations. For
presentation today I have smoothed them out severely so that neither the
highest peaks nor the lowest depressions now appear. These graphs speak for
themselves and require little comment or explanation (Fig. 1).

The first curve shows the number of printed pages in the Club’s publica-
tions which have heen used for taxonomy. It shows the feeble results of the
Club’s activities during its struggling first decade; the rapid rise of taxonomy
in the nineties, as Britton and Rusby came into action and as the BULLETIN
became a national rather than a local organ; the huge productivity in taxonomy
at the turn of the century when those active young men Britton, Small, and
Rydberg were at their best; and the gradual decrease in total taxonomic
matter in the last three decades as space became available in several new
publications. Since the curve is smoothed it does not show the peak of publica-
tion, which was 932 pages in 1906, nor the lowest point of the last half century,
which was 101 pages in 1926.

The second curve shows the percentage of total publication which has dealt
with taxonomy. It shows the almost exclusively taxonomic interests of the
membership in the early days of the Club, followed by twenty years of gradual
diversification ; a temporary rise over another twenty years, as the unparal-
leled productivity of New. York botanists overbalanced the generally growing
interest in morphology and physiology; a general period of decline during
the next thirty years, as the interests of the members became more diversified ;

38 LORREY A

and finally a rise in the proportion during the last decade, doubtless in response
to the general revival of interest in taxonomy.

The magnitude and importance of the Club’s contribution to the advance-
ment of taxonomy by means of its publication is, I am sure, realized and
appreciated by all taxonomists, and I trust that my figures have served to
make it clear to the non-taxonomic members of the audience.

Pages Pusiications oF THE TORREY Botawnicar CLus, 71870-7941,

5° realli tha ea ey ent | | | “- 00
| | | |
hae Sos roa iperrnl Be
| | | Number of pages of Taxonomy |
800 ! | t
\ ITN | | — — Percentage of space devoted
vy \| awl to Taxonom
700 SS SS ea Han \ | SS
tise | | Nadi / | | \\ | | |
600 rat =} |
gates idealyes Racglinn lose ge |
500 fea t | cesar 50
haga Neca a |
lakes | | er SS S\N
| AG
300 ; 30
| | | TN Sy
| i,
200 Y, 20
Cee | | | | |
| } | Note all curves smoothed |
| | } | | |

1930

Fic. 1

As a second and minor contribution I may mention the development of the
Torrey Club herbarium. Begun so long ago that I fail to find the date of its
inception, this herbarium grew very gradually through the donations of the
local members. Not long after the Museum Building of the Botanical Garden
was completed the herbarium was transferred to it, and continued to expand
through the yoluntary activity of interested local botanists and through the
collections of the Garden staff. The Club then presented the herbarium to
the Botanical Garden and it has since been maintained as a separate unit,
covering the area known as the Torrey Club range, which is roughly all the
territory within a hundred miles of New York, and illustrating the flowering
plants and ferns of this region by some 65,000 mounted specimens. The

GLEASON: CONTRIBUTIONS OF THE CLUB TO TAXONOMY 3°

herbarium may be consulted by any person interested in the local flora, which
is almost completely represented.

A botanical club, considered as a unit, can of course do no research, and
the Torrey Club has not employed taxonomists for research nor given grants
in support of it. Besides the two contributions to taxonomy which I have
already mentioned, is there any other way in which the Torrey Club can be or
has been of genuine service? There is a third way, which may not occur to
you immediately, in which the Club has been active, and through which,
measured by the extent and importance of the results, the Club has rendered
a highly valuable service, a service which has been partially outmoded by the
changed conditions of the twentieth century, but for which there is still an
opportunity and a demand. I refer to the encouragement and inspiration of
botanists. Botanists, like poets, are born, not made, but after birth they must
be developed. Today we have colleges and graduate schools for that purpose,
but such was scarcely the case in New York in the seventies and early eighties.
Even a formal education is not always sufficient. Probably every one of us
can look back to our earlier years and remember the inspiration which we
received from some one botanist, an inspiration which may have determined
us to become botanists rather than to enter some other profession.

Obviously, the professional botanists of New York today were not made
into botanists because of the influence of the Torrey Club, nor do they remain
botanists for that reason. Conditions were different sixty or seventy years
ago, when the death of Dr. Torrey left the Club without a leader and the
botanical interests of its members were kept alive largely through the encour-
agement of mutual contact, through the emulation of their fellow-members,
through the stimulation of new ideas, through the applause for the work
they accomplished.

There are some professions which can easily demand one’s full time, leaving
no opportunity for a hobby; there are some which offer excellent opportunities
for productive research to those who are so minded. There are still others in
which the prospect of large financial gain acts as a stimulus to continuous
work. Financial success, once it has been attained, is also apt to lead one to
devote his leisure time to the more fashionable forms of pleasure.

I shall cite to you five men who were trained and educated in a different
line, who earned their bread and butter in a different profession, whose interest
in botany was merely a young man’s hobby, but who maintained this interest
throughout their life and in two instances finally made it their life’s work.
One of these men had political advancement apparently within his reach,
but turned from it to enter botany at the bottom of the professional ladder. A
second had opportunity for research in a different subject. A third turned from

40 PPO RAR TE WEA

his original profession into botany before he was thirty. Two achieved financial
independence and still remained botanists by avocation.

Surely-there was a cause for this continued interest in plants, and I fail
to find any plausible cause other than the factor of encouragement and inspira-
tion received through the Torrey Botanical Club. Then, when you hear the
results achieved by these men, when you realize the part they have played
in the development of American taxonomy and in the provision of taxonomic
opportunity for others, you will agree that the most important contribution
yet made by the Torrey Club has been the inspiration and encouragement of
these men and of others whom I have not time to mention. The five are suff-
cient to demonstrate my point.

Eugene P. Bicknell, as a boy, was an amateur ornithologist and began
publishing in that subject at the early age of eighteen. As a man, he was a
banker. It was undoubtedly his membership in the Torrey Club and the stim-
ulus which he derived from it that gradually converted him into a clever
botanist. He was an exceedingly careful and discriminating observer of plants
in the field, and the bulk of his published work deals entirely with his field
studies. He was among the first to take his taxonomy into the field and to
base his conclusions primarily on his personal observations and only second-
arily on herbarium material. Do not understand from this statement that all
his taxonomic predecessors had been exclusively herbarium botanists ; nothing
would be farther from the truth. But, in general, they had formed their ideas
first in the herbarium and then substantiated them in the field, while Bicknell
- reversed the procedure.

His results were astonishing. Right here in the vicinity of New York, where
botanical work had been carried on for a century, he began to discover unde-
scribed species. Eastern botanists were surprised to learn, through his careful
field work, that there were more than one species of Helianthemum in the
vicinity. The common black snakeroot had always been referred to a single
species, or to a species and a variety, and Bicknell showed conclusively that
there were four. Scrophularia had held a single species in the eastern states,
and here he found a second. A grimonia had long contained only two accepted
species ; Bicknell’s careful field study showed several others. In rapid succes-
sion he turned his attention to other genera, Carex, Sisyrinchium, Lechiea,
Asarum, Teucrium, Rubus, Rosa, and various grasses, and in every case his
detailed and complete observations threw new light on their taxonomy. In
Rubus in particular, he early pointed out that the characters of the micro-
species of blackberries are of a different nature from those of the hawthorns,
and this observation, based on field study alone, is now being confirmed by
cytogenetics.

—

GLEASON: CONTRIBUTIONS OF THE CLUB TO TAXONOMY 41

In short, it was Bicknell, more than any other man of the period, who
returned taxonomy to the field and who re-opened the eastern states for
taxonomic research. In the great revival of taxonomy during the last quarter-
century, our own region has been found a fertile field for investigation. I do
not claim that Bicknell was directly responsible for this, but it is obvious that
he was followed, not preceded, by such similarly careful field men as Deam,
Stone, Wiegand, Marie-Victorin, and Fernald. The Torrey Club may well.
be proud that it had a part in this development through its encouragement and
support of the work of the banker, Eugene Pintard Bicknell.

The second man whom I shall mention was a successful lawyer, a promi-
nent judge in the New York courts, Addison Brown. He was a member of
the Torrey Club during the seventies, but being already established in his
profession he had less time and opportunity for field work. His botanical
work was chiefly centered on the collection of the various kinds of alien plants
which appeared on ballast dumps in the vicinity of New York City. His
few printed papers, published in the early volumes of the BULLETIN, show that
he collected many rare or unusual plants, some of them previously unknown
in America. His collecting stations are now mostly covered with buildings
and ballast-dumps are a thing of the past, but his specimens, conserved in the
herbarium of the New York Botanical Garden, show that his results were
accurately reported. Judge Brown’s contributions to botany were chiefly finan-
cial. It was he who assumed the financial responsibility for the publication
of Britton and Brown’s Illustrated Flora, without which the work could never
have been issued. I believe that I am correct in saying that no single book
ever did as much as this to revive and stimulate interest in the native flora
of the northeastern states and that his willingness to underwrite it derived
from his faith in Britton and his personal interest in plants, for both of which
the Torrey Club is responsible.

The third man was a geologist, who worked for a short time at mining and
then became a sanitary inspector for the City of New York. Interested in
politics, deeply concerned with all forms of civic improvement, he was soon
taking an active part in the affairs of the city and was appointed to several
city positions of increasing dignity and responsibility. In the middle of this
career he returned to science, which he had always followed as a hobby, entered
the graduate school, received his degree of doctor of philosophy, and became
one of the leading paleobotanists of America. Arthur Hollick’s name and
reputation are familiar to all of us and many of us remember him personally,
so that further comment is unnecessary.

The fourth man was also a geologist who, for some five years after the
completion of his work at Columbia College, was employed by the Geological
Survey of New Jersey. During this time he seldom missed a meeting of the

42 © TRARSE Vee

Torrey Botanical Club, and his interest in botany, increased and encouraged
by the Club, soon led to his determination to choose botany for his future
career. Accordingly he accepted a minor position at Columbia College, was
rapidly promoted to a professorship, and retired as professor emeritus at the
early age of thirty-seven. His name was Nathaniel Lord Britton, and his
retirement from the educational field was only to enable him to devote his
tireless energy to the development of the New York Botanical Garden. It was
his understanding and vision which led to the building of a scientific institu-
tion rather than a specialized park, to the accumulation of.a great herbarium
and a splendid taxonomic library, and through them to the provision of oppor-
tunity for taxonomic research by two score members of his staff, by some
hundreds of visiting taxonomists, and through the loan of herbarium material
by still more botanists in all parts of the world. In this place and before this
audience we do not need to dwell on the taxonomic achievements of Britton.
They are well known to all of us. But let us remember, as Britton himself
remembered, that to the Torrey Botanical Club he owed his botanical inspira-
tion and that to the Club he returned his thanks by his final generous provision
for its permanent endowment.

Fifth and last is a physician, Henry Hurd Rusby, whose name first appears
in the BULLETIN of the Torrey Club in 1878. So interested in botany was he
that even before he completed his medical education he had spent much time
collecting plants in the southwest, and soon after receiving his medical degree
he left for South America to explore for medicinal plants ; a search which was
successful, as we all know. This mixture of botany and medicine made of him
a pharmacognocist. During the remainder of his long life, 42 years of which
were spent as professor and dean at the New York College of Pharmacy, he
had every incentive to devote his energies entirely to pharmaceutical education
and the fight for pure food and drugs, in which he took a prominent part.
Without doubt, it was the enthusiasm which he drew from the Torrey Club
which led him to continue botany as his hobby and to devote to it every
possible minute which he could save from his regular work. Even in his last
decade, when failing eyesight made botanical work exceedingly difficult, he
continued to study his collections and to write short articles.

In 1887 Rusby had before hirn his extensive collections of South American
plants, largely made by himself but supplemented by many sheets from the
older Bolivian collectors Mandon and Bang. None of them was named; com-
parative material was scanty in the herbarium of Columbia College, and even
current literature was poorly represented in the Columbia library. So far as
North American botanists were concerned, South America was almost terra
incognita. Undismayed by the difficulty of the task, Rusby set to work on these
plants and also enlisted the aid of the rapidly rising young botanist, N. L. Brit-

Cin ON CONPRIBUGLONS @OnMmDaE CLUB IDO LAKONOMY 43

ton. Rusby made three later trips to South America and never lost his interest
in its flora. Neither did Britton, although he delegated most of the work to
others, returning to it personally only in his later years and especially after
his retirement in 1929.

These studies of the flora of South America grew and spread to other
American institutions and are primarily responsible for all our present interest
in South American botany. The important taxonomic work of Johnston and
Smith of the Arnold Arboretum, Moldenke of the New York Botanical
Garden, Killip of the National Herbarium, Pennell of the Philadelphia Acad-
emy of Sciences, Standley of the Field Museum and several others, have all
evolved directly or indirectly from the initial work of Rusby.

Rusby’s career as a taxonomist was peculiar. I fail to find that he ever
contributed to the general theory of classification, that he ever wrote a tax-
ononmic monograph, that he was ever a leader in taxonomic thought. But
Rusby was a two-fisted fighter, absolutely fearless of consequences to himself,
who fought adulterated food and impure drugs with the same intrepidity that
he faced the Amazonian jungles, who never admitted defeat and who seldom
was defeated. And here again I fail to find that he ever fought for a question-
able cause or for his own personal advantage. Instead he was a champion of
the right, as he understood it, and his understanding was correct.

Rusby was among the earliest to agitate for a botanical garden in the City
of New York and one of the leaders in the struggle for the necessary manda-
tory legislation at Albany. Later the directorship of the newly chartered garden
was in controversy and it was Rusby more than any other one person who
fought and worked to prevent the office from being merely another political
plum and to effect the appointment of N. L. Britton.

It has been my desire to express here my admiration and respect for one
of our former members, but my-words are too feeble for my thoughts. Henry
Hurd Rusby has gone from among us, but the results of his influence, his
energy, and his courage continue and widen from year to year.

Finally and in summary: The Torrey Botanical Club has not merely served
as a publishing agency, but it has also produced men, and these men, by their
additions to knowledge, by their provision of opportunity, by their influence
on modern thought, have been the chief contribution of the Club toward the
advancement of taxonomy. Let us hope that the Club will be equally useful
during the next seventy-five years.

THE New York BoranicaL GARDEN
New York, New York

VoL. 43 oe @VRARAE WON Jury 1943

Modern Taxonomy and Its Relation to Geography*

Henry K. SvENsSOoN

Taxonomy in the last seventy-five years has had increasingly close connec-
tions with geography, but the subject is so vast that only a small portion of the
field can be covered at this time. The most that can be done is to review some
of the geographical theories that have been in the light for two decades or more,
and with which we all are more or less familiar. These subjects are so inter-
twined that separate discussion of any of them is difficult and all of them are
but loose ends of the tangled thread that represents our fund of knowledge of
plant geography.

As to geographical location, we are practically astride the terminal glacial
moraine which runs the length of Long Island, and which was a collecting
ground for Asa Gray when he was associated with Torrey in New York. Much
ink has flowed on the subject of glaciation and its effect on plants since Gray
published his remarkable report on the similarities of the flora of eastern Asia
and eastern North America, in 1859. This date, which coincides with that of
the “Origin of Species,” was only eight years before the founding of the Torrey
Club, which can therefore be said to have occupied practically the whole
period of modern biology. Gray’s remarks were based on a collection by Charles
Wright, who is also well-known for his collections in Cuba and for those in his
own part of the Torrey Club Range, in Hartford, Connecticut.

As every taxonomist knows, the genera and even many species which we
find in our southern Appalachians are the same as those of the mountains of
western China and of Japan. The following quotations are from Gray’s paper,
Amer. Acad. Arts and Sci. Mem. 6: 1859: “The fundamental and most difh-
cult question remaining in natural history is here presented; the question
whether this actual geographic association of congeneric or other nearly rela-
ted species is primordial and therefore beyond all scientific explanation, or
whether even this may be to a certain extent a natural result. The only note-
worthy attempt at a scientific solution of the problem is that of Mr. Darwin and
Mr. Wallace,’ partially sketched in their short papers, ‘On the Tendency of
Species to Form Varieties; and on the Perpetuation of Varieties and Species
by Natural Means of Selection’ ” (p. 443).

“At length, as the post-tertiary opened, the glacier epoch came slowly on—
an extraordinary refrigeration of the northern hemisphere, in the course of ages
carrying glacial ice and arctic climate down nearly to the latitude of the Ohio.
The change was evidently so gradual that it did not destroy the temperate flora,

* Read at the 75th Anniversary Celebration of the Torrey Botanical Club at The New
York Botanical Garden, Tuesday, June 23, 1942.
- 1 Journ. Linn. Soc. (Zoology). 3: 45. 1858.
44

SVENSON: TAXONOMY IN RELATION TO GEOGRAPHY 45

at least not those enumerated above as existing species. These and their fel-
lows, or such as survived, must have been pushed on to lower latitudes as the
cold advanced . . ., portions of which, retreating up the mountains as the cli-
mate ameliorated and the ice receded, still scantily survive upon our highest
Alleghenies, and more abundantly upon the colder summits of the mountains
of New York and New England.”

“... perhaps the most interesting and most unexpected discovery of the
expedition is that of two strictly Eastern North American species of this order
[ Berberidaceae],—each the sole representative of the genus,—viz. Caulophyl-
lum thalictroides, and Diphylleia cymosa, of Michaux .. . are we to regard
them as the descendants of a common stock .. . or are we to suppose them in-
dependently originated in two such widely distant regions?” (p. 380).

“Smulacina (Majanthemum) bifolia extends around the world, but under
three pretty well marked geographical varieties :—the European, which extends
to eastern Siberia; the var. Kamtschatica, which replaces the former on the
Pacific Siberian coast, in Japan, and in North America west of the Rocky
Mountains; and the var. Canadensis, throughout all the northern part of this
country east of the Mississippi and the Rocky Mountains” (p. 414).

These quotations, it will be seen, are important from three points of view in
our modern taxonomy: 1) affirmation of the idea of evolution by the natural
selection of variations, 2) the negation of the bicentric origin of species, 3)
recognition of a holarctic Cretaceous flora of common origin, and its disruption
by the Glacial period.

And we arrive here at one of our first taxonomic difficulties. Shall these
geographic variants, which Gray showed to be of common origin, be classified

as a single species or shall they be segregated as separate species? This impor-
tant question we cannot decide. As Weatherby” has noted: “so long as we have
to rely on judgment at all, the accuracy and soundness of any taxonomic cate-
gory, definition or no definition, will be in direct proportion to the accuracy
and soundness of judgment of the individuals who apply it.” The pendulum
swings this way and that over periods of time. For example, the yellow lady’s
slipper (Cypripedium pubescens) of eastern United States has long passed as
distinct, but only recently Correll* has with some justification treated it as a
variety of the Eurasian Cypripedium Calceolus. The common brake of eastern
North America, long held as a separate species under the name Pteridiuim
latiusculum, has recently been returned by Tryon to its very old status ‘as a
subdivision of the wide-spread Pteridium aquilinum. And in Rhodora for this
very month of June we find the common water-plantain, which through later
years we have been patiently calling Alisma subcordata, blooming forth after

2 Rhodora 44: 160. 1942.
° Harvard Bot. Museum Leaflet 7: 1-18. 1938.

46 TORREYA

a fashion as a small-flowered variety of the Eurasian Alisma Plantago-aquatica.
Not only are species and their subdivisions the product of opinions of indi-
viduals, but the same is true of the limits of genera and of higher groups. Not
much is to be gained by a painful recital of the infinite variation of nomencla-
ture under present conditions; it is much more illuminating to review the geo-
graphic conditions which have made or should make a background for nomen-
clature.

This problem of glaciation in eastern North America has been ably treated
by Professor Fernald. He has shown that many species of restricted distribu-
tion in Western Newfoundland, in the Gaspé Peninsula of eastern Quebec, and
in some areas adjacent to the Great Lakes, are ancient plants (in contradistinc-
tion to Willis’ “Age and Area” hypothesis) that have persisted in places not
covered by the Wisconsin stage of the Pleistocene glaciation. The vegetation of
the glaciated area we may presume to have been obliterated during the ice age,
and since the deposits of the coastal plain are of comparatively recent origin
(chiefly marine) the uplands of the southern Appalachians and the Ozark
Mountains remain as areas from which the flora now inhabiting the coastal
plain has probably been derived. These various areas are shown in detail in
Fernald’s* recent work on the Virginia coastal plain.

Species which cover the three main areas (Appalachian uplands, glaciated
area, and coastal plain) often show marked divergences in structure in these
individual areas, and constitute geographic varieties, which if the variations
increased (according to the Darwinian interpretation mentioned in my open-
ing paragraphs) might become distinct species. The Appalachian plateaus still
harbor many species of the coastal plain, and from my own observations on the
vegetation of the barrens of Middle Tennessee, it seems probable that such
plants as Panicum meridionale, Rynchospora macrostachya, Scleria reticularis,
and Eleocharis microcarpa have moved into the Great Lakes area from the
siliceous uplands of Tennessee and Kentucky through Indiana, as we may infer
from isolated occurrences in the last named state. The bicentric range of
Lilaeopsis carolinensis in south-eastern United States and in the Argentina
region of South America is also shown. A similar disrupted distribution is
common in other groups, especially in the Cyperaceae, and is well shown in a
number of species of Eleocharis. Whatever may be the geographical explana-
tion, the problem of correlating published varieties and other subspecific units
in variable species with such bicentric ranges is well-nigh insuperable; it is
perhaps the most cogent argument for the non-recognition of varieties.

We may now turn attention to a recent publication of extraordinary interest
by J. C. Willis,? the author of “Age and Area.’ This work, entitled: “The

* Rhodora 42: 367. 1940.

> Cambridge University Press, England. 1940. 200 pp. Quotations by permission of The
Macmillan Company, publishers, U. S.

—

SVENSON: TAXONOMY IN RELATION TO GEOGRAPHY 47

Course of Evolution by Differentiation or Divergent Mutation,” is a negation
of evolution by the natural selection of variations as propounded by Darwin
and Wallace and affirmed by Asa Gray. The idea is not an original one, but is
based largely on what Guppy called “differentiation,” in his work on the vege-
tation of tropical islands. The eleventh chapter leads out, “Natural selection,
being a common phenomenon of everyday experience, has exercised such a fas-
cination that it has to a notable extent inhibited people from trying properly to
think out how a principle, whose essence is competition with partial escapes
into usually temporary success every now and then by improved adaptation, can
produce the ordered arrangement, taxonomy, and morphological or structural
uniformity with which we are familiar” (p. 103). Many, if not most or even
all, of the characters of distinction that mark families, sub-families, and even
smaller groups, are such that they can have no serious value upon the physio-
logical side which is the only one that matters from the point of view of natural
selection or gradual adaptation. These mutations are assumed by Willis to re-
quire long periods of time and to occur infrequently. “If one suppose a genus to
give off new species more or less in proportion to the area that it covers (which
again will be more or less in proportion to its age among its peers), it is clear
that all the offspring will carry a large proportion of the characters of the par-
ent, and that therefore while offspring arising near together will be most likely
closely to resemble one another, there is no reason why a close resemblance
should not arise with a wide geographical separation” (pp. 155, 156). “It will
commonly be found, in studying the distribution of the species of a genus, es-
pecially if it be of small or moderate size, that they are more densely congre-
gated toward the centre of the distribution of the genus, and fall off gradually
toward the edges, so that when one draws a line round the outermost localities
of each species one obtains a picture not unlike that which is called a contour
map by geographers ...”’ thus, Willis illustrates the distribution of the species
of Ranunculus found in New Zealand (pp. 149, 150). “Here one finds ‘wides’
(as I have called the species which have a dispersal outside the country in ques-
tion) occupying the whole area of the islands of New Zealand, and also reach-
ing eastwards to the Chatham Islands, 375 miles away .. . The endemics are
evidently crowded together rather south of the middle of the South Island,
whilst they fade out completely before the north end of the North Island is
reached ... The general impression that one gains from a map like this is that
the genus Ranunculus entered New Zealand probably from the south, and at
some place in the southern half of the South Island, where the incoming species
began giving rise to endemics, and on the average each species, wide or en-
demic, spread to the distance allowed by its age, and suitability to the conditions
with which it met.”

48 MEO RAR YAN

We now come to a region which has played a prominent part in taxonomy,
the Galapagos Islands and the adjacent coast of South America. These islands
were visited by Darwin in 1835, and upon the variations of birds and tortoises
from island to island, as well as upon the plants which were named by the
younger Hooker, were laid the foundations of evolution by geographic isola-
tion. The plants were briefly discussed by Darwin in the “Origin of Species”
(p. 349) : “Dr. Hooker has shown that in the Galapagos Islands the propor-
tional numbers of the different orders are very different from what they are
elsewhere. All such differences in number, and the absence of certain whole
groups of animals and plants, are generally accounted for by supposed differ-
ences in the physical conditions of the islands; but this explanation is not a
little doubtful. Facility of immigration seems to have been fully as important
as the nature of the conditions.

“Many remarkable little facts could be given with respect to the inhabitants
of oceanic islands. For instance, in certain islands not tenanted by a single
mammal, some of the endemic plants have beautifully hooked seeds; yet few
relations are most manifest than that hooks serve for the transportal of seeds
in the wool or fur of quadrupeds. But a hooked seed might be carried to an
island by other means; and the plant then becoming modified would form an
endemic species, still retaining its hooks, which would form a useless appendage
... trees growing on a continent, might, when established on an island, gain
an advantage over other herbaceous plants by growing taller and taller and
overtopping them. In this case, natural selection would tend to add to the
stature of the plant, to whatever order it belonged, and thus convert it into 2
bush and then into a tree.”

It is interesting to note in this connection that the only genus of plants now
recognized as endemic to the Galapagos Islands is Scalesia, which is bushy or
sometimes nearly herbaceous in the lower parts of the islands, but some species
become large trees where the moisture is more plentiful. Stewart in 1911 esti-
mated that 40 percent of the plants (varieties and forms being included in
the count), were endemic, but as in the case of the birds, the larger percent
of the endemic plants occur in a few groups. Many supposed endemics, further-
more, have been recently found on the desert coasts of Ecuador and northern
Peru; these areas have a climate strikingly similar to that of the Galapagos
Islands, and together with the islands seem to form a marked geographic
province. Taxonomic problems which vex the botanist have cropped up among
the ornithologists. For example, J. Huxley writes of Swarth (quoted by Gold-
schmidt in ‘‘The Material Basis of Evolution,” p. 209), “after classifying them
[the Galapagos finches] into five different genera with over thirty species and
subspecies, .. . it would be almost as logical to put them all in one genus and
species.” So far as the Galapagos are concerned the astounding extremes of

SVENSSON: TAXONOMY IN RELATION TO GEOGRAPHY 49

¢climate which occur in the same or adjacent localities in both islands and the
mainland may perhaps become as important to taxonomists as the question of
isolated land masses. Nor do these examples complete the difficulties of the
taxonomic picture. The coasts of Ecuador and Peru have been visited in a
desultory manner by botanists for over two centuries: Feuillee, Cavanilles.
Ruiz, Pavon, Humboldt & Bonpland, Hartweg, Andersson, Spruce, and
Weberbauer. New species were described bountifully, more frequently than
not without any references or comparisons with what had been described be-
fore. In this repect there is still much room for improvement in taxonomy.
This brings us to the last item, the question of taxonomy in respect to the
organism as a whole. A number of recent papers might be mentioned, but none,
it seems, quite comes up to the recent Memoir of the Torrey Club by Stebbins,
“Studies in the Cichorieae; Dubyaea and Soroseris, Endemics of the Sino-
Himalayan Region.” Such a treatment includes taxonomy, anatomy, cytology,
morphology of pollen grains, and probable phylogeny, especially in relation to
the geography of the species. If taxonomy in general were treated with such
care, many of our most distressing problems of nomenclature would vanish.

BROOKLYN BoTANIC GARDEN
Brooktyn, NEw York

VoL. 43 TOURS Yes Jury 1943

Some Economic Aspects of Taxonomy*

E. D. MERRILL

One dictionary definition of taxonomy is: “Classification; especially clas-
sification of animals and plants according to their natural relationships; also
the laws and principles of such classification.” Another, a bit longer is: “The
laws and principles of taxology, or their application to the classifying of objects
of natural history; that department of science which treats of classification ;
the practice of classification according to certain principles.” And in this same
dictionary taxology, a term I have never wittingly used, and which I shall
eschew, is defined as: “The science of arrangement or classification; what is
known of taxonomy.” Here I infer that the lexicographer responsible for the
definition of both taxonomy and taxology may have preferred the latter to the
former, but taxonomy, widely and universally used, will scarcely be replaced
by taxology, no matter what a lexicographer may prefer.

Under the first definition, including the laws and principles of classification,
one could wander far afield and become bogged down in discussions of the laws
of nomenclature for nomenclature cannot be disassociated with taxonomy, for
we must, of necessity, use names for the objects with which we are concerned.
However, I have no intention of thus widening the subject to include problems
of nomenclature and interpretations of the rules and regulations set up by inter-
national botanical congresses to govern the application of names, for such dis-
cussion would be endless.

This topic was assigned to me and is, perhaps, not one that I would have
chosen voluntarily. Thus I feel relatively little personal responsibility as to just
how I may develop the subject, realizing very fully that no two individuals
would treat it in a comparable manner. To limit the definition to “classification
according to natural relationships’ would be unwise, for in practice, while it 1s
fully realized that arrangement according to natural relationships is the objec-
tive that is always desirable, this is not always practicable. Often our reference
collections are totally inadequate, and we have to do the best that we can with
what is available. The result is that not infrequently we are obliged to utilize
characters of a more or less obvious nature, and not always those that indicate
the closest natural relationships between various groups, whether these be
major or minor categories. Again, we may utilize a combination of obvious
utilitarian characters associated with others that clearly indicate natural
affinities, in order to attain a certain objective.

As long as the learned world of the early European civilizations up to and

* Read at the 75th Anniversary Celebration of the Torrey Botanical Club at The New
York Botanical Garden, Tuesday, June 23, 1942.

; 50

MERRILL: ECONOMIC ASPECTS OF TAXONOMY 51

including the middle ages knew and utilized only a few hundred basic plant
species, botanical science and taxonomy was indeed a simple matter. In those
distant days a rough classification, as to major groups, as trees, shrubs, and
herbs sufficed. Species were designated by shorter or longer descriptive Greek
or Latin sentences. But even in these early days there was, here and there, the
beginnings of classification by obvious characters indicating varying degrees of
natural relationships. In the Europe of renaissance the pulse quickened. Up to
this time those who were at all concerned with plants and their utilization,
being scholastically minded, could think only in terms of the ancient Greek and
Latin masters. All attempted to refer their plants to those recognized and
named by the classical authors, particularly Dioscorides. In northern Europe,
with the invention of printing and the general advancement in learning, it
became evident that many of the species characteristic of this part of the con-
tinent were really different from those of the Mediterranean region. Once this
break came with classical traditions, progress was greatly accelerated, as evi-
denced by the masterful works of Fuchs, Brunfels, Bock, and others, for these
pioneers had returned to the actual study of plants as opposed to merely a study
of the classics. Following the epoch making discoveries of the pioneer Portu-
guese and Spanish navigators the small stream of botanical knowledge became
a flood.

Still for the most part the cumbersome system of designating species by
descriptive sentences prevailed and no radical change was made in nomencla-
ture until 1753, when Linnaeus promulgated his very simple and very obvious
binomial system. I say ‘‘very simple and very obvious” because it was so simple
and so practicable that one constantly wonders why it was not developed as a
system some centuries earlier. The idea of the genus had taken root at an earlier
date, and following Linnaeus’s innovation this radical departure in designating
plant species by a binomial, a generic and a specific name, quickly prevailed.
After all, in common everyday parlance the binomial system of designating
plants was widely used among the common people of many countries, but there
was a wide gulf between daily usage of the people and the learned world. Wit-
ness binomials in the common names of plants, such as white oak, red oak,
cork oak, burr oak, live oak, scrub oak, swamp oak, post oak, chestnut oak,
valley oak, holm oak, pin oak, water oak, willow oak; stone pine, sugar pine,
white pine, red pine, yellow pine, nut pine, Scots pine, Austrian pine, black
pine, loblolly pine, jack pine, and digger pine. This system of common names
as binomials is not modern, but is one of the most ancient things in many
languages, this usage being very widespread in the world at large, and among
primitive as well as among culturally advanced peoples.

But coupled with the Linnaean binomial system was his artificial system of
classification based essentially on the number of carpels and the number and

on
bo

TiO RE VY Awe

arrangement of the stamens. This was a very practicable system for arranging
genera as a matter of convenience and it dominated the field for somewhat
longer than the succeeding half century, although by the end of the eighteenth
century the handwriting was on the wall, and in the early part of the nineteenth
century the artificial system was generally replaced by the natural system of
classification with which we are familiar.

If the proposal of the binomial system by Linnaeus raised a mild storm
among those accustomed to the earlier much more cumbersome system of no-
menclature then in vogue, a storm that quickly subsided leaving the binomial
system universally established and accepted, the proposition to arrange the
genera in natural families raised a veritable hurricane among the devotees of
botany accustomed to the simple and convenient Linnaean system. This storm
raged for some decades and we of the present age have little conception of it.

In 1831, John Torrey published his American edition of Lindley’s
“Tntroduction to the Natural System of Botany.” He states in advertisement:
“Tn France, the natural or philosophical method has for many years past taken
the place of the artificial sexual system of Linnaeus, and recently by the
labours of Brown, Lindley, Hooker, Greville, and others, it has begun to be
employed in England and Scotland. .... I at once perceived that a desideratum
in British and American botany, long felt and lamented, was at length sup-
plied. It therefore occurred to me that I could not do a more acceptable service
to the friends and cultivators of Botanical Science in the United States, than by
preparing an American edition for the press forthwith. .... This is an epitome
of modern philosophical Botany, and will be found highly useful to those who
wish to obtain an accurate knowledge of the Natural Classification of the
Vegetable Kingdom.”

At this time all botanists in the United States, with the exception of
Rafinesque, were professed Linnaeists; there was no other system of classi-
fication as far as they were concerned. What happened? Consider Amos Eaton’s
statement of 1833.1 In speaking of Torrey’s edition of Lindley he wrote:

“Since Dr. Faustus first exhibited his printed bibles in the year 1463,
no book, probably, has excited such consternation and dismay as Dr. Torrey’s
edition of Lindley’s Introduction to the Natural System of Botany. And to
make the horrors of students, as well as of ordinary teachers still more
appalling, Dr. Torrey’s Catalogue of American Plants at the end of his Lind-
ley, was so singularly presented, that it would seem to indicate an awtul
catastrophe to all previous learning. To relieve all concerned, let me make
this pledge: Nothing new is presented either in the text or in the catalogue
[i.c., Eaton’s own Manual]. excepting what ought to have been discovered in
this progressive science, since the fifth edition of this Manual was printed; and

1 Eaton, A. Manual of Botany for North America, ed. 6, i-vi. 1833.

MERRILL: ECONOMIC ASPECTS OF TAXONOMY 53

not much real improvement has been added, as between the fourth and fifth
editions. . . . . As far as I have any influence I pledge it here, that the
embarrassing innovations of De Candolle and others are no possible use to the
ScIence Ol Botatnye sa An attempt is made in his Lindley to prove that the
Artificial method of Linnaeus is unnecessary. In doing this he proposes an
Artificial Method” of eleven pages. As those who have not read Torrey’s
Lindley will scarcely believe this unaccountable absurdity, they are requested
to examine, unbiased, that work between pages Ixvi and Ixxx of the introduc-
tion. This artificial system [artificial key to families] is said to lead to the
Natural Method..... The improvements upon Linnaeus, which have been
made, do not authorize any change in the science of Botany other than mere
additions and corrections. ... .

This caustic critique of the natural system of classification is eliminated
from the seventh (1836) and eighth (1840) editions of Eaton’s “Manual,” and
in these, although he adhered to the Linnaean artificial system of classifica-
tion, he so far relented as to include an epitome of the natural system. If,
however, one needs a good illustration of a closed mind, here we have it, and
this statement is made in all due regard to Eaton’s remarkable accomplish-
ments® although it is only fair to explain that in botany Eaton never claimed
originality. He states* that in the field of botany he never aspired to be anything
above that of a teacher, translator, and compiler. It should be noted that Eaton
italicized his characterization of botany as a progressive science, yet at the same
time insisted that the suggested improvements on the Linnaean system did not
authorize any changes in the science of botany other than mere additions and
corrections! This is an ultra-conservative, nay, even a reactionary attitude.

McAllister, p. 235, quotes from John Torrey’s letter of November 2, 1833,
to L. D. von Schweinitz giving his reaction to edition 6 of Eaton’s “Manual”:
“This time Torrey was more effusive (italics mine) in his praise of the
Manual when he wrote to his friend De Schweinitz “Have you seen the 6th edn.
of Eaton’s Manual of Botany ? .... 1 began to read the preface in a bookstore
the other day & it seemed to be a most remarkable performance.’ In view of
the circumstances one wonders if the term “effusive” is the corect one, for in

2 Eaton apparently wrote this very hurriedly, for this statement regarding an artificial
method is an error. What is presented is an artificial analysis of the orders in the form
of a key to the classes (Vasculares, Cellulares), subclasses (Exogenae or dicotyledonous
plants, and Endogenae or monocotyledonous plants), tribes (Angiospermae, Gymnosper-
mae, Petaloideae and Glumceae), and to the families under each division and subdivision,
these, as to limits (but naturally not as to sequence as at present understood) much the
same as they stand today. Torrey’s “singularly presented” catalogue is merely an arrange-
ment of the genera of North, American plants by families under the natural system!

3 McAllister, Ethel M. Amos Eaton. Scientist and Educator, i-xiii, 1-587, illus. 1941.
* Manual. ed. 7. iv. 1936.

54 ; AV OMRGROT NG

the same letter Torrey also says’ that he had scarcely seen more than the covers
of the book and that he was interrupted before he had finished the first page;
and this first page begins with Eaton’s castigation of Torrey, my quoted
passage: “Since Dr. Faustus first exhibited his printed bibles in the year 1463,
no book has, probably, excited such consternation and dismay as Dr. Torrey’s
edition -of Lindley’s Introduction to the Natural System of Botany.” I am
afraid that the dear lady didn’t read this preface, for under the circumstances
Torrey’s statement to De Schweinitz can only be interpreted as sarcastic
and ironic, as far as a gentle soul like John Torrey could be ironic and sar-
castic, certainly not as “effusive” praise! The relationships between Eaton and
Torrey had their ups and downs. Clearly we do not have to confine our reading
to the opinions of modern botanists to learn just how certain individuals judge
their contemporaries, for throughout botanical history individuals have not
hesitated to say just what they thought about the work of this or that author.
In the constant quibbles that one notes in taxonomic literature one is reminded
of a remark ascribed to President Lowell when some acute problem regarding
the interrelationships of certain prima donnas among Harvard botanists
needed to be settled: “What is it about the pretty little flowers that makes the |
botanists quarrel so much among themselves ?”

Within a decade or two from the time that Eaton castigated Torrey for his
progressiveness, the Linnaean system of classification was entirely outmoded
and abandoned, and was replaced by the natural system that he so violently
condemned. Eaton, the non-progressive botanist 1s, as a botanist, only a vague
memory among the devotees of this science today. But Torrey, who was the
subject of his scorn, forged steadily ahead to become the outstanding American
botanist of his time ; and this organization, the Torrey Botanical Club, the old-
est botanical association in America, today celebrating the seventy-fifth anni-
versary of its establishment, honors John Torrey’s name, and its founders
incidentally honored the organization itself, in the selection of its name, a
perpetual reminder of the services rendered by this outstanding individual and
botanist. Had Torrey been another Eaton, clearly there never would have been
a Torrey Botanical Club. :

Because of the vast number of organisms that the naturalist must deal with
as to species, to say nothing of higher categories such as genera and families,
it is clear that it is impossible to arrange large groups m any lineal arrange-
ment that will show all natural relationships. This is particularly true of the
major groups. We may follow the Bentham and Hooker system for convenience,
treating in sequence first the dicotyledonous plants, then the gymnosperms, and
then the monocotyledonous groups, although this is a very unnatural arrange-
ment because the gymnosperms are infinitely more pfimitive, among the flow-

siMemy Tor Bot Clabes 290, 1921

MERRILL: ECONOMIC ASPECTS OF TAXONOMY 55

ering plants, than the dicotyledons and the monocotyledons. Or we may select
to follow the Endlicher system as developed by Engler and Prantl, treating the
gymnosperms first, then the monocotyledons and finally the dicotyledons ; or
we may decide with Wettstein and others, that the dicotyledons should be
placed before the monocotyledons if the system is to be a natural one, in
accordance with various lines of evidence as to the comparative times of
development of these last two groups.

It is inevitable that when a proposed system becomes very widely used,
like that of Bentham and Hooker, or that of Engler and Prantl, it will become
more or less fixed, partly from the weight of authority, partly because of con-
venience and for comparative purposes. We may all realize that the Engler
and Prantl system of arranging families, in some respects is far from a natural
one, and that radical changes are indicated, particularly in reference to the posi-
tion, in sequence, of such families as the Magnoliaceae, Ranunculaceae, Ber-
beridaceae, etc., which seem clearly to be much more primitive than the
Amentales, for example. System after system may be proposed, but relatively
few of these will, from the very nature of things, become widely accepted as
to the sequence of arrangement of major groups, partly from inertia on the part
of working botanists, partly because it is always desirable to be able to make
direct comparisons with the work of others, and partly because one is never
sure as to just when some morphologist may discover evidence that upsets
all previously proposed systems and sets up another “improved” one. It all
comes down to the simple fact that within the plant kingdom, when one is deal-
ing with such groups as natural families, it is impossible to make any lineal
arrangement that will show all relationships and inter-relationships, for devel-
opment and differentiation has not followed a straight line from a lower to a
higher group, but in many cases it has been divergent, and, we may suspect,
reversions have played their part. To indicate natural relationships we must
construct variously branched “‘trees” to show origins and relationships as well
as historical sequences ; but in a book we must hew pretty closely to the straight
line, whether we are dealing with a series of families in a system of classi-
fication, or whether we are dealing in terms of a simple manual for field use,
for one page follows another from beginning to end.

Again, we must always keep in mind that the objects with which we are
dealing are variable; that our accumulated knowledge constantly increases ;
that a system that we might set up today, on the basis of the available data, may
be outmoded a few years hence when more comprehensive collections, and
when a more intensive study of obscure details, perhaps supplemented by
anatomic, cytogenetic, genetic, historic, and geographic data, become avail-
able. This comment applies more to the problem of species and their inter-
relationships than it does to larger categories such as genera and families. All

56 OTRAS At

active systematists are familiar with these factors from their own daily work.
As examples, I may cite my own experience. In 1904, I hopefully prepared a
key to the 21 then known Philippine species of Medinilla, not realizing what
changes would be necessary within a few years, for less than twenty years
later, about 125 species of this genus had been described from or accredited
to the Philippines. In 1900 there were actually known from the Philippines
only 13 species of the Pandanaceae, Freycinetia with 7 species, and Pandanus
with 6, of which only one was definitely understood and could be placed in ref-
erence to other described species of this genus, five described by Blanco appear-
ing in all botanical literature as species ignotae or species dubiae. Twenty-five
years later not only had all of Blanco’s “unknown” species been placed, but the
total for the family stood at 93 species, Freycinetia 45, Pandanus 47, and
Sararanga 1. This is what has happened in family after family and genus after
genus within the present century as comprehensive collections have been
assembled from the botanically little known parts of the world such as China,
the Philippines, Malaysia outside of Java and to a certain degree the Malay
Peninsula, Siam, Indo-China, tropical Africa and tropical America. What is
the reaction of local taxonomists, working on a restricted flora, the con-
stituent elements of which are well known, in reference to such a work as that
of Schlechter® in which no less than 1153 new species of orchids are described in
one work, and these all from German New Guinea? The area of German New
Guinea is 68,500 square miles, and for comparison that of New York State
is slightly less than 50,000 square miles. Incidentally, approximately 2500 new
species of orchids have been described from the Island of New Guinea since
1900. These cited examples merely represent a few that demonstrate the
acceleration of what happened within the present century as various parts of
the world were opened up to botanical exploration. What happened in various
parts of the world happened in the United States when the West was opened up
by exploration, and still later when a respectable body of ‘local botanists
developed in the West. This is, in part, the basis of the break between Asa
Gray and E. L. Greene, for Greene was on the ground and was intimately
acquainted with the local flora of California; I say “in part’’ because there was
also an entirely different concept between the two as to what constituted a
species.

It will be a long time yet, at our present rate of progress—which may be
greatly slowed down in the coming years—before the imperfectly known
regions mentioned above may be considered to be even reasonably well
explored. Until this end is attained all treatments of all large groups that
have representatives growing in these vast and only partly explored areas can

® Schlechter, R. Die Orchideen von Deutsch-Neu-Guinea. Repert. Sp. Nov. Beih.
1: i-lxvi. 1-1079. 1911-14.

MERRILL: ECONOMIC ASPECTS OF TAXONOMY

Ur
N

be considered only as tentative. We do the best that we can with what we have
at hand, and optimistically hope for the best. One closing example. In 1800,
about 65 species of Ficus were more or less definitely known from the entire
world. In 1801 Willdenow' described four new species and rather naively
remarked: ““Je ne doute pas que dans le climats chauds il n’existe encore
plusieurs especes de figuiers encore inconnues,” little realizing that before the
year 1940, a total of approximately 2,400 binomials would actually be pro-
posed in this Brogningnagian genus—God forbid that these 2,400 binomials
represent 2,400 distinct species, but the number of valid ones is very great,
certainly approaching 2,000, even without splitting hairs on specific differences.
If any taxonomist is looking for new worlds to conquer, I recommend that he
undertake a monographic treatment of this vast assemblage.

In citing the above examples of the rapid increase in the numbers of pro-
posed species in certain genera, far be it for me even to suggest that the actual
naming and describing of new species is an end in itself, or if there is anything
difficult about the art. As a matter of fact it 1s a very easy and simple matter
to name and describe a species as new, it isn’t so easy to determine whether or
not the particular form in hand has been named and described by some earlier
botanist or whether it actually constitutes a sufficiently distinct entity to be
considered worthy of consideration as a species; to say nothing about
macrospecies or microspecies, nor even to mention subspecies, variety, sub-
variety, form, proles, or any other category that has been suggested, but never
too well defined, to indicate minor entities. With the myriads of forms with
which we must deal we must have names. The competent monographer fol-
lows and either embalms our possible error by recognizing a species as valid,
or sinks it into synonymy; and if the latter happens then at some future
date some other monographer may reinstate it with the chances that in the
interim some other optimistic taxonomist may have renamed and redescribed
the same form under a new name in his confidence that a published reduction
is always a reduction, which, perhaps unfortunately, is not always the case.

The special properties of a very high percentage of our thousands of species
of economic plants, whether utilized for food, for medicine, for fibers or for
any other purposes were originally discovered by empirical processes and by
observation rather than by direct and deliberate investigations. This is the
history of most plant species of economic importance whether it be the lowly
bean used for food, or the insignificant looking Ephedra sinica now extensively
utilized in the practice of medicine. Although this Ephedra has been utilized
by the Chinese for many centuries it is only within the present century that it
was definitely demonstrated that its curative principle ephedrine is really of

* Willdenow, C. L. Determination de quelques nouvelles espéces de Figuier, et cbserva-
tions générales sur ce genre. Mem. Acad. Sci. [Berlin] 1801: 91-104. f. 2-5. 1801.

58 PRORREN AL

distinct value in the treatment of asthma and various diseases of the nasal
passages. Through taxonomy, however, a realization of the relationships of
plants, we find what may be an important lead. If Ephedra sinica yields
ephedrine, isn’t it possible or even probable that other species of the same
genus may yield the same curative agent? Thus a pharmacological investiga-
tion of all species of Ephedra might be indicated, for the sole natural source
of Ephedra sinica is northern China, although other species of the genus occur
in various parts of Asia, Europe, and North America. It is admitted, now that
ephedrine has been synthesized, that further work on representatives of this
particular genus may scarcely be worthwhile, but the case serves to illustrate
the problem of botanical analogy.

Take the case of chaulmoogra oil, now extensively and successfully used
in the treatment of leprosy. For centuries this oil was used in India for the
treatment of leprosy and various skin diseases. For nearly a hundred years
the situation was confused because the plant named by Roxburgh as Chaul-
moogra odorata Roxb., but never actually described by him, was supposed to
be the species that yielded the effective drug; yet the seeds of Roxburgh’s
species, later described as Gynocardia odorata R. Br., when investigated, were
shown to contain no active curative principle. It was not until 1900 that Sir
George Watt cleared up the confusion and determined the botanical source of
the true chaulmoogra seeds as Taraktogenos Kurzu King = Hydnocarpus
Kursii Warb. Rock,* who has discussed this subject, states that it is quite
probable that not only seeds of this species but also those of H. castaneus
Hook. f£. & Th. and other species of Taraktogenos and Hydnocarpus, as yet
undescribed, are sources of the chaulmoogra oil of commerce. The botanical
confusion that prevailed for a hundred years unquestionably retarded a critical
and serious investigation of chaulmoogra oil as a remedy for leprosy. It is only
within the present century that this cure has come into its own.

Intrigued by the problem of analogy and suspecting that the seeds of some
of the Philippine species of Hydnocarpus might contain the same curative prin-
ciples as the true chaulmoogra oil, I was instrumental in fostering an investi- _
gation of those Philippine species that were available, including Hydnocarpus
Alcalae C. DC., H. subfalcata Merr., H. Woodii Merr., and H. Hutchinsonii
Merr. Various studies were made in the Bureau of Science culminating in
1928, when Messrs. Perkins and Cruz® investigated the oils of ten species
including four from the Philippines and Borneo, and found that in these four ~
species the oil was very similar in chemical composition to commercial chaul-

8 Rock, J. F. The Chaulmoogra tree and some related species: A survey conducted
in Siam, Burma, Assam, and Bengal. U. S. Dept. Agr. Bull. 1057: 1-29. ¢. 1-16. 1922.

® Perkins, G. A. and Cruz, A. O. A comparative analytical study of various oils in
the chaulmoogra group. Philip. Jour. Sci. 23: 543-569. ¢. 1. 1928.

MERRILL: ECONOMIC ASPECTS OF TAXONOMY 59

moogra oil except that Hydnocarpus Alcalae C. DC. contains a very large
amount of chaulmoogric acid and little or no hydnocarpic acid. The total per-
centage of oil varied from a minimum of 11 percent to a maximum of 39 per-
cent. Now as far as known none of the Philippine and Bornean species was
utilized for any purposes by the native population. They were, of course,
unknown to the small technical public outside of the very few botanists, and
it is an interesting commentary to note that as to the Bornean Hydnocarpus
Woodu Merr. trees were actually found to be growing within the limits of the
leper colony on Sandakan Harbor; a remedy actually at hand, but previously
unknown, and its potentialities hence unrealized.

In the latest treatment of this group’? Taraktogenos Kurz is reduced to
Hydnocarpus Gaertn. and a total of forty species are recognized. Not more
than one-fourth of these species have been investigated from a pharmaceutical
standpoint ; and yet from what is known of the properties of those that have
been investigated it is safe to assume that the seeds of most of the species of
the genus will be found to yield the same curative principles as are found in
the true chaulmoogra oil.

Thus from analogy, working from a Burmese species, the curative principles
in its seeds being known, investigations extend to the seeds of the Philippine
and Bornean species of the same genus, Hydnocarpus, with potentially impor-
tant economic results. These examples will suffice to demonstrate what has
been done in special cases, and by analogy we may expect that in the future
similar investigations will be extended to very many species that have hitherto
never been considered as even worthy of investigation; but in a reasonable
percentage of cases we may definitely assume that these species, as yet unknown
and unappreciated from an economic standpoint, will be shown to produce
needed and otherwise unattainable products. Here the tempo increases under
the pressure of necessity brought about by war conditions in reference to sup-
plies of rubber, quinine, and various other products for which, in the past, we
have depended largely on Asia and Malaysia for our supply ; and our economy
and even way of life was increasingly geared to various imported basic prod-
ucts which now are unobtainable elsewhere. Now new sources must be devel-
oped, if not from the same species so successfully developed in the specialized
agriculture of certain parts of the Old World (even although in some cases
based on native American plants, such as Hevea and Cinchona), then from
others that yield similar products. It is in this specialized field of potential
substitute plants that may yield important products that we now lack, that the
trained and experienced taxonomist can render, and is rendering, fundamentally

10 Sleumer, H. Monographic der Gattung Hydnocarpus Gaertner nebst Beschreibung

und Anatomie der Friichte und Samen ihrer pharmakognostisch wichtigen Arten (Chaul-
mugra). Bot. Jahrb. 69: 1-94. t. 1-4. 1938.

60 T_ORIRE YOR

basic services. [t is this type of individual who knows his plants and who
knows plant relationships who can serve to great advantage, for his accumu-
lated store of special knowledge cannot be matched by those botanists trained
and experienced in other fields remote from that of taxonomy and systematic
botany. Let us hope that those charged with selection for super-specialized
services such as those indicated in this field of botanical analogy, will select
wisely and well. After all there is much truth in the popular conception of what
a botanist is—an individual who knows and can name plants; yet the vastly
higher percentage of our professional botanists have almost no knowledge and
less experience in this specialized field of taxonomy, and many of them have no
interest in it. They are for the most part specialists in totally different branches
under the all-inclusive term botany, for in our times the term botanist covers
not only the taxonomist and systematist, but also the fields of morphology,
physiology, ecology, cyto-genetics, cytology, histology and various other
subdivisions ; the numerous devotees to these subdivisions of botany are all
“botanists” in spite of the popular definition cited above. A very high percent-
age of them would be utterly lost were they to be assigned to special problems in
this distinctly complicated field of botanical analogy.

Within the field of medicine or pharmacology, here is a simple illustrative
case. The European Digitalis purpurea Linn, is the source of an important drug,
digitalin, and we have generally depended on Europe for our supply. With
these supplies now cut off by the war, local sources must be developed. I
have no idea of how extensively the plant is now cultivated in the northern
United States, but Fernald, on the basis of his own extensive field knowledge,
calls attention to the fact that the species is not only thoroughly established in
certain parts of Newfoundland, but that in places it is dominant and a
veritable pest; a source of supply that only needs to be tapped if there be
need to build up our dwindling stocks, and an indication that certain parts
of Newfoundland are ideally adapted to the actual cultivation of the species on a
large scale if this be needed.

It is clear to all taxonomists and all systematic botanists, that in spite of
the imperfections in our current system of naming and describing plant
species, and in spite of the distinctly Rafinesquian character of the work of
certain individual botanists who can see differences where tangible differences
scarcely exist, that taxonomy and the accurate identification of plants is basic
to a proper understanding of myriads of problems in the general field of
economic botany, pharmacology, agriculture, plant breeding, piant pathology,
genetics, forestry, morphology, physiology, and many other fields into which
plant science or botany sensu latiore has been subdivided. We have little
patience with the investigator, no matter what his problem may be, who ignores
this basic problem of accurate identification of the material with which he

MERRILL: ECONOMIC ASPECTS OF TAXONOMY 61

deals. Obviously if one deals with misidentified material his findings may
prove to be valueless, for future investigators will find it difficult if not impos-
sible to check his results. There are too many errors in botanical literature due
to this lack of critical consideration of this simple basic problem, and much
time, and some space in our technical periodicals, has been wasted due to the
ignorance or the blind faith of investigators, or those who have stimulated
research on a particular subject, who have not considered it to be either essen-
tial or even worthwhile to check, or to have some competent taxonomist
check, the identity of the plant utilized to prove this or that conclusion. Here
is a horrible example:

In 1902 there was published in one of our leading botanical magazines a
paper on the morphology of the flower and embryo of Spiraea that admirably
illustrates the importance of accurate identification. The investigator worked
with material representing a single species, the plant widely known among
horticulturists under the erroneous name of “Spiraea japonica.” Far from
being a representative of Spiraea or even of the family Rosaceae this plant is
Astilbe japonica A. Gray of the Saxifragaceae. The author completed his
detailed study without even suspecting that he was dealing with a misidenti-
fied plant, from which we may assume that he could not have done much
bibliographic research as the differences between Astilbe and Spiraca are
remarkable. Is this blind faith in a labelled growing specimen or sheer careless-
ness or ignorance on the part of those who suggested and supervised the
work and thus victimized an innocent graduate student who had faith in the
knowledge of his preceptors? The net result was to discredit the student, for
about all he got out of it was some training and experience in laboratory
technique, discredit to the periodical in which the article appeared, and, may
we hope, some discredit on those who sponsored the investigation. It is a
classical example of how not to elucidate a morphological problem, for the net
result merely served to stimulate the glee of the lowly taxonomists who, as a
group, are thoroughly satiated with the “holier than thou” attitude of some of
our colleagues in the laboratory aspects of botany. I am much less charitable
than was Rehder who called attention to the error.

What do we taxonomists think, when we observe in a physiological paper a
tabulation of species whose seeds will not germinate until after they are sub-
jected to freezing temperatures and note the strictly tropical Carica Papaya
listed in this category? True, pawpaw and papaya are common names of
Carica Papaya but pawpaw is also the common name of our entirely different
northern Asimina triloba Dun. We can only assume that the seeds of Asimina
were what this investigator had, for Carica is a plant entirely intolerant to
freezing conditions. All of which merely illustrates that we should not put our
trust wholly in the currently used common names of plants. After all, “What is

62 TORRE VY #

in a name, a rose by any other name would smell as sweet’’ but in cases like
these, one is reminded of an expression used by one of the characters in that
intriguing comedy, “You can't take it with you” when he was expressing his
opinion of the dancing ability of another character in the play.

In this part of the discussion J am rapidly approaching a category recently
discussed in the daily press. Under date of May 18, it is reported from
Raleigh, North Carolina, that some years ago the Daughters of the American
Revolution planted, with elaborate ceremony, a little tree purported to be an
offspring of the “Continental Elm” at Cambridge, Massachusetts, under which
George Washington is supposed to have taken command of the Continental
Army in 1775. They even kept a box of earth taken from around the roots of
the parent tree for use in christening the “elm” when it grew up. The little
“elm” has grown up and is now blooming; but it is a cherry tree and not an
elm at all. Assuming that the young tree that was planted was provided by
some nurseryman this merely proves that nurserymen and _ horticulturists.
can make mistakes just as botanists do, but is this any reason why a botanist
making a really serious study of a plant problem should accept without ques-
tion as to its correctness, a commonly used but erroneous horticultural name,
or should determine what binomial he should use merely by looking up a
common name?

One closing example, that of the investigator who had laboriously dug up
and intensively studied the root tips of Tilia in one of our large collections, and
could not understand the discrepancies between the chromosome counts of the
root tips and of the branchlets taken from the same trees in a number of cases.
It was only after the study had been completed, but fortunately not published,
that he learned that many of these species of Tilia were grafted, the roots
representing an entirely different species from the growing tree. Thus for cer-
tain types of investigations we cannot even trust the living plants without
knowing something about their history.

I have above referred to the fact that during the many centuries Europe
was dependent on its own economy, its inhabitants utilized only a relatively
few plant species ; a few hundred important ones at most. As various parts of the
world were opened up within the few centuries following the expansion of
the European colonizing nations the number of species utilized rapidly
increased ; and this tempo of increase continues unabated. In 1853, Linnaeus
recognized 5,950 species of plants in all groups for the entire world, while he
and his immediate followers estimated that there might be as many as 10,000
species of plants, in all groups, in the world. The estimate had been increased
to 30,000 known species by 1820, and 50,000 indicated as probable for the entire
world. By the middle of the century the estimate of known species was 93,000.

Within the present century about 265,000 new binomials have been pub-

MERRILL: ECONOMIC ASPECTS OF TAXONOMY 63

lished for the flowering plants and vascular cryptogams alone, of which about
194,000 represent hopefully proposed new species, the remainder shuffles or
transfers from one generic name to another. The yearly average for the higher
groups alone is now approximately 6,500 as new binomials, of which about
4,750 represent proposed new species. This is the record of the twentieth
century to date. The total number of binomials published from 1753 to 1942
is in the neighborhood of 750,000 for the higher groups of plants alone, and to
this must be added those published for the cellular cryptogams; our grand
total should be in excess of 1,000,000.

As to the total number of distinct and more or less “known” species, who
shall say? Jones has briefly discussed this matter’ calling attention to the
remarkable discrepancies that occur in recent texts, with a spread in the esti-
mates of from 133,000 (Uphof’s estimate of 1910) to 175,000 for the angio-
sperms alone, and concludes that the total for all known groups is in the
neighborhood of 335,000. Because of various complications that it is unneces-
sary to discuss here, I suppose that we may conclude that one guess is as
good as another; but knowing something about synonymy; something about
the limiting factors in the geographic distribution of individual species ; some-
thing about more or less universally distributed species ; something about the
extraordinary richness of tropical floras ; something about the remarkable local
endemism in various tropical areas; something about the high percentage of
novelties that are found in all new collections from hitherto inadequately
explored areas; something about those regions that, within the past four
decades, have been particularly rich in the crop of new species—my guess
is pretty close to that of Jones, and that the total number of reasonably valid
described species in all groups is well in excess of 300,000. Even if the num-
ber of valid species should be only half this total, what scientist, no matter what
his field, would even have the temerity to suggest that we can get along with-
out taxonomy and nomenclature?

In this discussion I have deliberately been discursive rather than specific.
One could cite case after case of the applications of taxonomy to various scien-
tific and economic problems, but a few will serve to bring out the points at
issue. Besides those mentioned above in my discussion of botanical analogies
we may list the problem of the Citrus relatives ; the case of Coffea arabica Linn.
versus Hemuleia vastatrix Berk.; Berberis versus wheat rust; the Pinus-Ribes
complex in reference to the blister rust of the white pines; the little problem
of special strains in such lowly organisms as the yeasts and the fungi when
these organisms are basic to certain industrial processes—the list would be
unending, for no agricultural crop exists in which problems of plant breeding,
of protection against fungus diseases and insect pests do not exist. Many prob-

1 Jones, G. Science IT. 84: 243. 1941.

64 fh OR RE WA

lems Lave been solved, but many more are still with us, 2nd new ones develop
from year to year. With all due regard to the qualifications and accomplish-—
ments of the specialists in the various fields concerned, I maintain that the
better equipped the investigator is in basic taxonomic knowledge, the better
is he fitted to work on his special problems. This does not mean that all
botanists should be taxonomists, but it does mean that all specialists and ail
laboratory botanists should realize the importance of accurate identification,
the implication of botanical analogies, and that they should appreciate the
facilities outside of their own fields that are available in specialized institutions
in various parts of the country. We will go much further with reasonable
cooperation than we will by maintaining a pigeon-hole type of specialization.

There should be no real antagonisms between the devotees of various
aspects of botanical science, for the inter-relationships are close—much closer
than some of our specialists realize. We are all laborers in the same vineyard,
and our objective is progress; progress in pure science as well as in the eco-
nomic aspects of the subject as a whole. To those representatives of the
laboratory school of botany who are hypercritical regarding taxonomists and
systematists, I would call attention to the fact that progressive taxonomists are
now taking advantage of the findings of their associates in other fields includ-
ing the histologists, pollen experts, geneticists, cytologists, ecologists, and
entirely outside of the biological field invoking the aid of geologists, hydro-
graphers, geographers and others in their attempt to solve certain prodlems of
plant relationships.

This very organization that this week celebrates the seventy-fifth anni-
versary of its establishment was founded by individuals whose fields of interest
were essentially field botany, taxonomy and systematics. It has evolved, during
the course of years into a national organization and has wisely and progres-
sively widened its activities, yet the unifying idea that maintains it is still that
of its founders who were interested in plants and who knew plants as they grew
in nature rather than merely as laboratory subjects. I repeat what I have
written before: “It has been fashionable in some quarters in modern times
to decry both the importance and the value of systematic botany. Because of its
vitality, its human interest, its practical bearing on other phases of plant
science, and on our everyday life, one suspects that some of its critics have
lacked the breadth of view of leaders in science, and have been misguided in ~
criticizing that which they did not fully understand.”

Let us take the broader view, live and let live, keep our respective houses
in order, avoid egregious blunders, and attain a realization of the fact that after
all there is a unity in plant science in spite of its diversity, and that the entire
field is interlaced with the binding bonds of system and order; and this is
taxonomy.

HARVARD UNIVERSITY
‘CAMBRIDGE, MASSACHUSETTS

Vow. 43 ARO IR IR 1D YON Jury 1943

The Importance of Taxonomic Studies of the Fungi*

FRANK D. Kern

The naming and classifying of living organisms has been going on for
centuries. It has been well said that “a large part of our thinking about living
things is bound up with some system of classification.” Another writer has
pointed out the fact that we depend much upon classification in our general
experiences. “It is the innate propensity of active minds,” he says, “to form
species, 1.¢., successively to make distinctions, to point out similarities, and then
to assemble the things that are alike into their kinds. It applies to everything
from chemical elements to college fraternities.”

The recognition of the need of names for plants dates from the days of
Pliny, the Roman naturalist, and Dioscorides, the Greek physician, in the first
century of the Christian era. Plants could not be discussed without names.
They could be named, however, without classification. They could be classi-
fied, also, without a conception of phylogeny. In other words, nomenclature
deals with names which may or may not be arranged according to a system
of classification; and classification deals with groups which may or may not
indicate relationships. Many biologists, on the other hand, attempt to arrange
groups on a basis of similarities, which they believe to be expressions of actual
relationships. It is of particular interest today to note that the modern
development of these aspects of botanical science has been made during the
years since the founding of this Club. The first real progress in working out a
universal system of nomenclature was made at an International Botanical
Congress in Paris in 1867. A natural system of classification, although early
recognized as desirable, has made its most progress since the theory of evolu-
tion provided a basis for phylogenetic interpretations. Darwin’s Origin of
Species, just a few years earlier, furnished the evolutionary concepts which
soon became so significant in taxonomy.

Even a cursory examination of some of the early attempts to classify the
fungi is sufficient to reveal that the results were most general in nature.
Bauhin, in the days of the “herbals” purported to bring together all the plants
known to him and to all those who preceded him (Pinax Theatri Botanici,
1623). The concept of the genus as a group of species had not then become
definitely established. In the group which he called Fungus were included 81

* Read at the 75th Anniversary Celebration of the Torrey Botanical Club at The New
York Botanical Garden, Tuesday, June 23, 1942. Contrjbution from the Department of
Botany, The Pennsylvania State College, No. 137. Publication authorized on July 6, 1943
as paper No. 1185 in the Journal Series of the Pennsylvania Agricultural Experiment
Station. :

66 AD (GYD NZ IAN

species which are now distributed to at least nine families. Tournefourt, in the
latter part of the 17th century, made a considerable contribution to the genus
concept. He recognized six genera of fungi and one of lichens. Dillenius and
Vaillant added some genera and the latter published illustrations which were
a real contribution to the study of the fungi. He maintained the genus Fungus
in which were included most of the forms of the family Agaricaceae.

The foremost pre-Linnaean student of the fungi was Micheli. By the time
of the publication of his “Nova plantera genera” in 1729 the microscope had
become a working-aid and he made use of it. His work was excellent for the
time. It included consideration of the genera of flowering plants, ferns, mosses,
lichens, algae, and fungi. Both large and small forms of fungi were given con-
sideration. He germinated and grew spores of the larger fungi and observed
both mycelium and sporophores.

The early workers who studied the microfungi under the microscope rather
naturally tried to interpret them in the light of their knowledge of the parts of
flowering plants. In the case of the bread-molds the sporangia seemed like
little fruiting pods containing seeds. By analogy rust spores were similarly
interpreted although the situation there was not so easily demonstrated as with
the molds. In 1807 DeCandolle, referring to the spores of Uromyces and
Uredo, said that “with a microscope this powder seems composed of ovoid or
globular spores ... . filled with many small grains that are considered spores.”
He thought that a teliospore might contain at least 100 such “spores.” This
interpretation prevailed among such workers as Fries, Léveillé, and the
Tulasne brothers, and persisted until the time of De Bary in the middle of the
19th century.

Linnaeus set himself the task of bringing together in his “Species
Plantarum” (1753) all the known species of the plant world. He included the
fungi in his class Cryptogamia but it cannot be said that he advanced the
knowledge of them to any appreciable extent.

The first author to make a distinct advance in the classification of the fungi
after the beginning of binomial nomenclature was Persoon. In a paper
published in 1794 (Neuer Versuch einer Sytematischen Eintheilung der
Schwamme, Romer’s Neues Mag. Bot. 1: 63-128) he recognized 77 genera
of fungi, which he placed in two classes: Angiothecitum and Gymnothecium.
The three genera of rusts, which were included, were the first rust genera to be
established after the solitary rust genus of Micheli 65 years before. Several
authors of important works during the first quarter of the nineteenth century
followed Persoon’s classification in the main. Among these were Schumacher,
Rebentish, Albertini and Schweinitz, De Candolle, and Brongniart. During
the same period Link brought out a new classification which was accepted
wholly or in part by Schlechtendal, S. F. Gray, and Wallroth.

KERN: TAXONOMY OF THE FUNGI 67

During the middle of the nineteenth century great contributions to the
knowledge of the larger fungi were made by Elias Fries. He had “not only a
poor opinion of the parasitic fungi but an antiquated conception of their
nature.” In his third volume of “Systema Mycologicum” (1832) he used the
name Hypodermi to include the rusts, smuts, and some other fungi and
characterized them as having “No proper vegetative body ; sporidia originating
from the metamorphose of the cellular structure of living plants: an inferior
kind of fungi.” Nevertheless the work of Fries which extended over more than
a half a century gave a great impetus to the study of fungi. His prestige was so
great that there were many who accepted his leadership. Among these may be
mentioned Endlicher, Léveillé, Corda, Rabenhorst, Strauss, Berkeley, and
Cooke. Most of these authors made changes in the arrangement of the genera.
Corda’s extensive publication (Icones Fungorum) is notable not only for its
contribution to the knowledge of the structure of the larger fungi but also for its
advances regarding hundreds of the microfungi.

During the first three quarters of the nineteenth century new species
were being recognized and named from all parts of the world. The descrip-
tions appeared in journals, reports, and books many of which were not widely
circulated. It is little wonder that investigators soon found it difficult to know
whether or not a species under consideration was already described and
named. It may be well said that this condition still exists. Thus it came about
that species were named and renamed from several to many times. Little was
known of the distribution of the fungi and workers in one region had no way
of knowing of the probability of the existence elsewhere of the species which
they were studying. Conceptions of the probable cosmopolitan distribution of
the fungi were necessarily slow in developing. Many efforts were directed
toward bringing together all species known to occur in certain regions or
countries without attempts to determine their wider distribution. The flora-
type of publication became common, especially in the European countries.
Rabenhorst’s “Kryptogamen Flora” of Germany, Austria, and Switzerland
is a good example. Many other floras could be cited. These publications were
valuable but they did not solve the problem for the workers who were located
away from the European centers of mycological activity.

The assertion that many mycologists actually were deterred “from describ-
ing supposedly new species for fear of duplication’”’ will doubtless not meet with
credulity. An important step toward overcoming this situation was the plan
for the “Sylloge Fungorum” inaugurated by Saccardo in 1882. The first
volume appeared in that year. The effect was an immediate stimulation of sys-
tematic mycological activity. This great work developed into twenty-five
volumes, the last appearing in 1931. During this period mycological journals

68 TORRE Yrs

were established in various countries and taxonomic work with the fungi went
forward at a rapid rate.

Thus far we have given consideration chiefly to the describing, naming and
classifying of the many and varied forms. The earlier workers naturally were
concerned with these phases of study. It should not be concluded, however,
that there were not some, even among the early workers, who were intrigued
with the possibilities of studying the development and life-histories of the
forms with which they worked. There were suggestions that relationships
might exist between different forms which were found in close association.
The impress left by De Bary on this phase of mycological work is well known. °
He began his work about the middle of the nineteenth century and the type
of investigation which it stimulated has continued up to the present. He found
time to work not only with fungi but also with algae, myxomycetes, bacteria,
and higher plants. It is said that no less than 68 workers, afterwards distin-
guished in science, studied under him at Strassburg. According to Erwin F.
Smith, “His work and that of his students put plant pathology on a new
foundation, and he also, undoubtedly had much influence on human and ani-
mal pathology, since his very successful infection experiments with fungi on
plants suggested many things to those who were trying to determine the cause
of human and animal plagues.” Yet we must agree that the primary interest
of De Bary was in morphology rather than in pathology.

Using a good microscope and employing micro-chemical reagents De Bary
made important advances in the knowledge of spores, infection, and mycelia.
His cultural demonstration of heteroecism in Puccinia graminis, with proof
that the aecidium on barberry was a stage in the life-cycle of wheat rust is well
known. These results were announced in 1865. This work, and more which
followed, ushered in a new phase of mycological endeavor. It is significant that
he began these investigations not out of pure scientific interest, but in order
to settle controversies between agriculturists and botanists regarding the rela-
tion between smuts and rusts and diseases. Agriculturists thought them to be
the causes of disease while botanists were inclined to regard them as products
of disease. De Bary had himself resisted the suggestion of a possible alternation
of generations which required an alternation of hosts plants. When his experi-
ments led to that conclusion, his naive statement that “one comes around, per-
haps, in a way, to the ancient opinion according to which rusted wheat would
be infected by the rust of barberry” is most interesting. His experiences should
be heartening to many present-day investigators who are required to work on
projects which are economic and agricultural in nature. Out of such problems
may arise basic scientific discoveries as in the case of De Bary.

The next epoch in the study of the fungi after De Bary was ushered in by
the study of the nucleus and its behavior. This gave a new direction to the

KERN: TAXONOMY OF THE FUNGI 69

study of fungi. As life-histories were important for taxonomic considerations
so nuclear developments were eventually recognized as having a bearing on
taxonomy. The application of cytological methods to the study of life-histories
in the fungi began with the work of Dangeard in 1894 and was soon under
way on a large scale. Other early workers in this field were Poirault, Sappin-
‘Trouffy, Maire, Harper, Blackman, and Christman. It was soon evident that
the nature of sexual reproduction in the fungi was of great value in determin-
ing relationships. We are indebted to such a host of investigators that it is
impossible to mention them by name. Notable studies have been made in the
Phycomycetes, Ascomycetes, Ustilaginales, Uredinales, and higher Basidio-
mycetes. In the last few years genetical studies have been made and highly
important results are in the making.

Our account would not be complete if we did not make some reference to
the possibility that the classification of the future may have a physiological
basis. Much headway toward such a goal has been made by Mez and his asso-
ciates. Many of you are familiar with the fact that Mez, using serological
methods, has constructed a family tree of plants which corroborates in a
remarkable manner the older tree based on morphological characters. Seifriz
refers to this work in a recent book (The Physiology of Plants, 1938) with the
remark, “It is of great significance to the field of evolution and phylogenetic
relationship that a purely chemical basis of classification should so well sup-
port a purely anatomical one.” Seifriz points out that the relationships between
plants established thus far by serology hold well for families, not so well for
genera, and not at all for species. He believes, however, that this is due to a
lack of delicacy in technique. He is of the opinion species differences in proteins
must also exist.

Our historical sketch which began with the early attempts to classify fungi
led us rather inevitably to some consideration of morphological, cytological,
genetical, and physiological studies. Certainly we must agree that knowledge
gained in all these fields is essential for progress in taxonomy. E. A. Bessey in
1939 (A Textbook of Mycology) refers to the present-day activity of sys-
tematic mycologists and points out that, “Life histories are being studied in
all groups, the sexual relations are being scrutinized from the lowest to the
highest fungi and genetical studies are revealing results somewhat parallef,
but on a vastly smaller scale as yet, to those attained by the study of Zea mays
and Drosophila.” “As never before,” says Bessey, “is a knowledge of fungi
themselves so necessary.” Obviously right conceptions of fungi must be based
upon many facts, and wrong conceptions can easily be the result of partial facts,
and of ideas derived from other plants which may be inapplicable and mis-
leading.

We have referred to the contribution which Darwin’s theory of evolution

70 AN OIRIRDS YS se

made to biological classification. Phylogeny soon became the fundamental basis
for classificatory endeavor. So far as the fungi are concerned we should not
overlook the influence of the work of Hofmeister in 1851 on the bryophytes
and pteridophytes. The recognition of an alternation of generations in these
groups had its effect on studies of the algae and fungi.

Every student who has taken a course in general botany is familiar with
the system of classification which places the algae and fungi together in the
division Thallophyta. We have no thought of attempting to reach any con-
clusions about this broad question of the taxonomic disposition of the fungi.
Whether the fungi are to be regarded as one of two subdivisions of the Thallo-
phyta, the algae being the other, depends upon the origin of the fungi. We say
this in spite of a recent assertion that the taxonomist “‘is not interested in the
origin, but in the character of his plants.” On the origin of the fungi, G. M.
Smith, in his “Cryptogamic Botany,” Vol. I, “Algae and Fungi’ (1938)
writes, “This is highly controversial and opinion is divided as to whether
they arose from the protozoa or whether they had either a monophyletic or
polyphyletic origin among the algae. If they arose from protozoa, they should
be put in one or more divisions coordinate in rank with the various algal
divisions ; if they arose from the algae, they should be placed as classes of one
or more of the algal divisions.”

Smith reviews the algal and the protozoan theories of the origin of the
fungi and concludes that “it seems more probable that the fungi evolved from
protozoa rather than from algae.” He bases his conclusion largely on metabolism
and the type of flagellation in the Phycomycetes. There are some algal groups
in which there occur chlorophyll-less forms which are so similar morpho-
logically that they cannot be regarded as distinct from the green forms. It is
pointed out that these saprophytic and parasitic algae accumulate reserve
carbohydrates as starch just as do the green algae. In contrast the Phycomy-
cetes are reported generally to accumulate carbohydrates as glycogen but
never as starch. The zoospores and gametes of the green algae are never uni-
flagellate whereas the motile cells of certain Phycomycetes are regularly uni-
flagellate. It is admitted that the question of the origin of the Ascomycetes is a
more difficult one. The similarity in the sex organs, and the structures developed
subsequent to fertilization, in the Ascomycetes and in the red algae are strik-
ing and have caused many workers to assume a relationship between these
groups. Smith argues that these distinctive reproductive structures may have
evolved along independent phyletic lines. He thinks the Ascomycetes had
their origin in the Phycomycetes and that the Basidiomycetes arose by modi-
fication from the Ascomycetes. In his classification he rejects the Thallophyta
as a division of the plant kingdom and in its place substitutes nine divisions,
of which the Myxothallophyta, or slime molds, constitute one and the

KERN: TAXONOMY OF THE FUNGI 71

Eumycetae, or true fungi, constitute another. The other seven divisions
include the algae. “Abandonment of the Algae as a subdivision of the plant
kingdom,” says Smith, “does not mean that the word alga must be abandoned.”
He believes that we can still use the term alga for designating simple green
plants that have an independent mode of nutrition. We might add that we
will likewise continue to use the term fungus although attempts to define it
lead to difficulties. |

Bessey in his “Textbook of Mycology” has attempted a definition of the
term fungi that would not commit the definer to any system of classification.
We quote: “Fungi are chlorophyll-less thallophytic organisms typically con-
sisting of coenocytic or cellular filaments, but including also encysted or
amoeboid one-celled organisms which reproduce by some type of motile or
non-motile spore; excluding the Bacteria and such chlorophyll-less organ-
isms, which, by their structure, are with definiteness assignable to recognized
orders of algae.’’ Bessey is of the opinion that the Mycetozoa are not related
to the fungi; are not, indeed, plants. There are those who believe that the fungi
should not be regarded as belonging to the Plant Kingdom. Herbert F.
Copeland in a comparatively recent paper (Quarterly Review of Biology,
December, 1938) has presented evidence and argument “to the effect that
organisms can be arranged, naturally, and more conveniently than in the
past, in four Kingdoms as follows” :

Kingdom 1. Monera (Bacteria and Blue-green Algae)

Kingdom 2. Protista (Protozoa, Diatoms, Red and Brown Algae, Slimemolds, and

Fungi)
Kingdom 3. Plantae (Green Algae, Liverworts and Mosses, Ferns and Allies, Seed

plants )
Kingdom. 4. Animalia (Metazoa)

To those who have been accustomed to thinking that all living organisms
must be either plants or animals the recognition of two new groups as King-
doms may seem revolutionary. It is true, however, that the line between lower
plants and lower animals has always been a difficult one to draw. It must
be admitted that nomenclatorially there are difficulties in placing together in
the Kingdom Protista organisms which have been previously in two different
Kingdoms. The original proposal for a Kingdom to be called Protista was made
by Haeckel in his ‘‘Generelle Morphologie” in 1866. He also established the
group Monera but included it in Protista. According to Copeland other
authors have expressed the opinion that the Monera should be treated as a
separate Kingdom.

The comments presented here relative to the origin of the fungi form a
very inadequate picture of the discussions and arguments that exist in the
writings of many investigators. We have wished merely to call attention to

12 DORREVS

the fact that there is no general agreement as to whether the fungi are
monophyletic or polyphletic in origin or whether they have descended from
the algae or from the protozoa. The algal theory appears to have been advocated
by A. Braun in 1847, and was accepted by Cohn (1854), Pringsheim (1858),
and Sachs (1874). De Bary in 1881 objected to the method of intercalating
the iungi among the algae saying it led to an orderly arrangement of species
but not to a natural system. The suggestion that the fungi arose from the
protozoa is credited to Cornu (1872), and was developed by Gobi (1885)
and Dangeard (1886). Atkinson (1907) was in favor of deriving the lower
fungi from ancestral unicellular organisms, but was uncertain whether they
were colorless or chlorophyll bearing. He was, however, certain that their
origin was monophyletic. The algal origin of fungi was supported by Stras-
burger and C. E. Bessey. Gauman (1925) presented the view that all true
fungi were derived from the green algae in monophyletic line; he believes the
lower Chytridiales (his class Archimycetes) along with the Myxomycetes may
have arisen from the colorless Flagellatae. He does not regard either of these
groups as fungi. Martin (Bot. Gaz. 93: 421-435, 1932) has “suggested that the
fungi be regarded as a phylum which has not definitely developed into either
plants or animals, but may be grouped with the former as a matter of con-
venience, and in accordance with custom.” He rejects the assumption that
all living organisms are descended from a single primitive cell and points
out that the assumption that life may have originated more than once and
in different forms is more in accord with what we know of living organisms.

Clements and Shear (Genera of Fungi, 1931) enunciate a basic prin-
ciple: “that the fungi do not constitute a natural group, and that all the
phyletic lines lead sooner or later to holophytic origins.” It should be noted
that although they say they are not dealing with a natural group yet they claim
to have approximated a natural system in several respects in their book. They
believe that there is but one natural system and they maintain that any
approach to it must be the result of the work of many minds. After their
admonition that it is more or less inexact, even though convenient, to con-
nect the name of an individual to any particular arrangement, one wonders
whether he should not tear up his manuscript and begin anew. Clements and
Shear do not agree that cytology can be the final arbiter on questions of origin
and relationship among the fungi. They make a plea for experimentation “on
the largest and broadest scale possible, in both field and laboratory.”

This review which is concerned with the taxonomy of the fungi must pro-
vide reference to the specialists who publish papers or monographs on certain
groups. Sometimes such authors are called experts. I like the way one writer
who says he is no expert disposes of this matter. He says, “The standard
taxonomic revision is the work of an expert in the group concerned; it cites

KERN: TAXONOMY OF THE FUNGI 73

all the present literature; it is received with respectful interest (never with
complete acquiescence) by the author’s fellow experts in the same group,
and is more or less annoying to others who have to take it into account, as
requiring revision of familiar ideas of the limits of groups and the applica-
tion of names.” The parenthetical phrase is not mine; it is in the original.

As with other groups of living organisms the fungi have had their
devotees. Crowds of them have advanced to the expert stage. It is- impossible
to name them or to evaluate their contributions. They must be treated
generically, as it were. The writer has thought it worth while to try to present
some of the problems which such workers encounter. By this is meant not so
much the problems inherent in taxonomic studies but rather the wider limita-
tions which often operate to check individual progress and to break the con-
tinuity of advances for which a groundwork may have been well established.
The difficulties which are to be discussed are not necessarily peculiar to sys-
tematic mycology. Taxonomic work in general as well as in mycology, has
a checkered history. Its advances through the centuries have been piecemeal.
Perhaps it will always be thus, and deploring the fact may not only be in
vain but may not be fitting. |

It seems likely that we must depend largely upon institutions to furnish
the support for taxonomic mycology. Of course there have been numerous
individuals who have done their work chiefly or wholly without institutional
support. In this country we have only to think of such men as L. D. von
Schweinitz, J. B. Ellis, C. E. Fairman, J. J. Davis, and Elam Bartholomew,
to realize the debt we owe to individuals, and great credit is due them.

Even where universities, colleges, or other institutions or governmental
agencies are involved it is still true that the ambition, industry, and perseverance
of individuals are largely responsible for the advances that have been made. In
these later days we have been hearing a good deal about institutional research.
So far as taxonomic work with the fungi is concerned we delieve that an
analysis would show that research in this line is mostly due to individual
prosecution rather than to institutional initiation. It may happen that an
institution will make an effort to continue the type of research that has been
inaugurated and successfully carried on by one of its staff members and will
then refer to the program as an institutional program. More ofteri it happens
that a real leader appears and develops successfully a line of work which is
supported (more or less) during his years of activity but which is dropped
by the institution afterwards. Such instances indicate the correctness of the
conclusion that there is often no such thing as an institutional program. There
are, of course, exceptions but we feel safe in saying that the exceptions prove
the rule rather than make it. We have inserted the parenthetical phrase—
more or less—because we are sure that institutional support even when

74 DEO VRIR Na

forthcoming during the height of the program is often more apparent than
real. Certainly it is true that many of our productive mycologists have had to
earn their “bread and butter” with teaching and routine duties and have
had left only a small percentage of their time and efforts for the kind of work
which they were so well qualified to pursue.

Someone may well ask why these difficulties are raised in connection
with taxonomic research when they exist in so many lines of research activity.
There are several reasons for doing so. The source materials for taxonomic
research are in large part not commercial commodities. They consist of rare
books, separates, indexes, illustrations and specimens which are accumulated
only with time, patience, correspondence, and exploration. When such col-
lections have finally been put together in an institution they should be used
by more than one generation of workers in that institution. Or if that is not
possible some method should be worked out by which they become available
to succeeding investigators in other institutions. There are now in existence
some collections of microfungi where spore measurements and drawings
accompany literally hundreds of specimens. Such aids are indispensable for
taxonomic studies and when available not only save the time necessary to
duplicate them elsewhere but help to prevent errors and misconceptions.
There are also herbaria of fleshy fungi where great accumulations of photo-
graphs, drawings, and notes make them of the utmost importance to other
workers. This is not a plea for the centralization of mycological taxonomy.
It is rather to call attention to the fact that enormous resources are fre-
quently accumulated and then not used nor made available for use. Since our
modern concepts fix the application of names by types rather than by descrip-_
tions it is a fair question whether type specimens should ever be personal
or institutional property. The difficulties may seem insurmountable but this
may not be the case. Surely we will make no progress until the workers them-
selves reach a keener appreciation of the situation. .

There are other factors which bear on the progress of taxonomic work
with the fungi. Even though a staff member may have the ability and enthu-
siasm to carry on work of this sort it may be, as previously indicated, difficult
for him to obtain the full cooperation of his institution. Projects which have
more evident economic aspects have always elicited more favor with adminis-
trative officials in our agricultural institutions. This is true in spite of the
obvious relation of taxonomic studies of the fungi to many phases of plant
pathology. It is easy to comprehend why this attitude prevailed in the early
days of the agricultural experiment stations but it is not so easy to see why
the value of fundamental work of this sort should not eventually come to be
recognized more generally. In very recent times approval of agricultural
projects depends upon evidence that results are likely to be of direct benefit

KERN: TAXONOMY OF THE FUNGI 72

to farmers. And again, even though there may be institutional approval so far
as the time of the worker is concerned, it is often difficult to secure the
maintenance support which is essential. For a project requiring special
apparatus, machinery, glassware, and chemicals, it is usually not difficult to
secure funds. But to secure funds for the purchase of specimens, photographs,
particular books, separates, periodicals, indexes, and exploration it may be
difficult or well-nigh impossible. It is generally conceded that a research worker
is not expected to get along with the equipment and supplies which are in
general stock but is entitled to special expenditures for his project. Not so
with library facilities. He may be expected to get along with what the institu-
tional library provides. He may of course compete for more than his share
of the general library funds but this is not always satisfactory even if partially
successful. The use of research funds for special library facilities is much less
common than for special material equipment. The problem of publication is a
closely related one. Monographic treatises are often expensive to publish and
the demand for them may be slight and slow. The fact that publication is diff-
cult tends to discourage this type of work.

A few weeks ago I received a letter from a former associate in which he
said, “I notice, with much interest, in the last issue of Science, that you are to
have a part in the ‘Symposium on Taxonomy,’ June 23, in connection with the
Seventy-fifth Anniversary Celebration of the Torrey Botanical Club... . I
assume that you will speak for the fungi.” Of course. Whether I have said, or
still can say, anything which he would have me say is another matter. I assume
that he expected me to make some reference to the problem of nomenclature
and it seems impossible to close this discussion without bringing up this
vexatious topic.

I propose to make comments of a general nature and to confine them to
two aspects of the nomenclatorial situation: (1) on getting rules, and (2)
on getting them into effect.

It is generally conceded that “Natural history can make no progress with-
out a regular system of nomenclature, which is recognized and used by the
great majority of naturalists in all countries.” This is a quotation of the first
article of the International Rules of Botanical Nomenclature; the italics are
mine. The necessity of establishing international rules to govern the applica-
tion of names of plants has been recognized by botanists for many years. But
it is easier to recognize the problem than to solve it. The world well knows
the difficulties of securing unanimity of action on any matters calling for
international consideration.

One of the chief difficulties is to get together a group, the personnel
of which is truly representative of the science and at the same time really
international in standing. Institutions and governments have been willing to

76 TORREYA

designate individuals as representatives to botanical congresses but for the
most part they have been unwilling, or thought it unwise, to contribute toward
the expense of attendance. The final assembly has been made up, therefore, not
necessarily of those best qualified but of those individuals who have been willing
to finance a trip in order to take part in the proceedings. The departments of
our national government sometimes send “official delegates” to international
congresses but they usually place restrictions on the activities of such delegates.
I hope I am giving away no secret when I say that an employee of our federal
government told me when we were in attendance at an International Botanical
Congress that he was instructed before leaving this country that he might
take part in the discussions but was not allowed to vote on the questions com-
ing before the section on nomenclature. The conclusion seems to be justified
that the advancement of this phase of natural history, of the greatest importance
to mankind, has been too dependent upon voluntary contributions of the
workers themselves.

It is also generally conceded that rules of nomenclature should not be
arbitrary and that they cannot be imposed by authority—at least not by the
authority of the makers of the rules. As an alternative the framers of the rules
say, “They must be simple and founded on considerations clear and forcible
enough for everyone to comprehend and be disposed to accept.’’ Such a state-
ment was made in the Rules as published in 1912 which were adopted in
1905 (Vienna) and supplemented in 1910 (Brussels). Perhaps rules of
nomenclature are like a plant which grows slowly and requires a period of
development before it comes to maturity. I do not know how many people
did not comprehend the International Rules of Vienna and Brussels but I do
know that in the following years many were disposed not to accept. There were
individuals and groups of individuals who deplored the fact that certain
fundamental principles of a basic nature in which they believed were not
incorporated. They felt that once they accepted a code without these principles
the chances for amendment would not be good. I have in mind chiefly the
“type-concept” which was not a part of the original code. Reference to a more
or less minor feature may serve to illustrate difficulties regarding adoption.
The Vienna code provided that “On and after January 1, 1908, the publica-
tion of names of new groups of recent plants will be valid only when they are
accompanied by a Latin diagnosis.”” Again I do not know how many names
have since been published which are invalid, but I do recall taking part in a
business session of a certain mycological society, at least 25 years after the
Latin deadline, whén the matter before the house was whether that rule
should be enforced in its official journal.

It seems fair to say that cordial agreement was reached at the Cambridge
Congress in 1930 on most of the disputed nomenclatorial problems and that

KERN: TAXONOMY OF THE FUNGI

MI
“SI

the disposition to accept International rules was improved thereafter. Not long
ago I was criticized by a colleague for such a conservative statement. He wanted
me to say that these rules are, and have been for some time, actually in effect.
Again it may be time which settles many problems. At any rate, it was in
1940 that the Secretary of the United States Department of Agriculture
formally approved a recommendation of the Department Committee on Plant
Names “to put the Department, botanically speaking, under the International
Rules of Nomenclature.” To me it is interesting that it took ten years for
this department to come to an action making these rules official for “publica-
tions, reports, and correspondence involving scientific plant names.” Perhaps
one might be pardoned for calling attention to the anomaly of an agency finally
finding it expedient to subscribe to the acts of an organization which it failed
officially to aid. It is also interesting to note that two years after the official
order they are still going through an adjustment period in getting nomen-
clatorial usage realigned according to International rules. When it becomes
necessary to drop the name Ustilago hordei which, according to old usage,
has been applied to the covered smut of barley and to take up the same name,
according to International Rules, for Joose smut of the same host it is little
wonder that the workers talk about confusion. Personally, I believe that the
confusion will be only temporary and that the advantage of getting on a
world-usage basis will more than outweigh the disadvantages. It is desirable
to avoid changes in names as far as possible, but changes cannot be entirely
avoided if the rules of nomenclature are to put in order the old names as
well as to be a guide for the creation of new names. There are those who
believe that the procedure embodied in the present system of nomenclature
leaves too much to expediency and personal preference and do not rest
sufficiently upon foundamental principles. It has been pointed out that “there
is no guarantee—if, indeed, there is any hope—that the system which may be
adopted today will be accepted by the next generation.” No, there is no
guarantee that anything man devises will continue—not even democracy. We
must not, however, look upon this or any other problem in such a futile
manner. There are difficulties, to be sure, but they are not insurmountable.
We are told in the Torrey Botanical Club Announcement and Field Schedule
for 1942, “It is understood that there will be no mutilation of species at this
session.” That being the case, this seems to be the proper place to bring tls
discussion to an end.

THe PENNSYLVANIA STATE COLLEGE
STATE COLLEGE, PENNSYLVANIA

Ser EE

Vor. 43 TRORREYA Jury 1943

AG TIYVAUDIESJOP Bib Gus
January TO May 1943

January 5. ANNUAL MEETING.

The annual dinner meeting of the Torrey Botanical Club was held at the Men’s
Faculty Club, Columbia University, at 6:45 p.m. The President, Dr. C. Stuart Gager,
presided, with 82 members and friends present. After the dinner the minutes of the
preceding meeting were approved. The reports of the Treasurer, of the Chairman
oi the Field Committee, and oi the Editor oi Torreya were distributed in mimeo-
graphed form, and the combined report of the Editor and the Bibliographer was read
by Dr. Matzke. These reports were accepted on a motion by Dr. Karling.

Dr. Gager addressed a few remarks to the Club and then announced that the following
list oi officers had been elected ior the year 1943:

President: William J. Robbins

1st Vice-President: Fred J. Seaver

2nd Vice-President: Lela V. Barton

Corresponding Secretary: Edwin B. Matzke

Recording Secretary: Honor M. Hollinghurst

Treasurer: W. Gordon Whaley

Editor: Harold W. Rickett

Bibliographer : Lazella Schwarten

Business Manager: Michael Levine

Members of the Council: Charles A. Berger, Clyde Chandler, Albert E. Hitchcock,

Roger P. Wodehouse
Delegate to the Council of the N. Y. Academy oi Sciences: Bernard O. Dodge
Representative on the Board of Managers oi the N. Y. Botanical Garden: Henry
A. Gleason

Representatives on the Council of the American Association for the Advancement

of Science: John H. Barnhart, Albert F. Blakeslee

Dr. Matzke then conducted a “Botanical Information Please” quiz with a board of
experts comprised of Drs. Gager, Graves, Karling, Robbins, and Zimmerman, aug-
mented at times by the guests at large. The meeting adjourned at 9:15 p.m.

Honor M. HoLt_incHurRST, RECORDING SECRETARY.

January 20. MEETING aT THE NEw York Botanical GARDEN.

The meeting was called to’ order at 3:30 p.m. by the President, Dr. William J.
Robbins. Attendance 25. The minutes of the preceding meeting were approved. The
following new members were elected: 15 to Annual membership, 3 to Associate mem-
bership; 2 transfers to Annual membership and 4 transfers to Associate membership
were approved. The resignations of 21 Annual and oi 4 Associate members were
accepted with regret. :

A letter was read concerning the preservation of High Tor. Dr. Robbins suggested
that a letter be sent to the sponsors of this movement, expressing the interest of the
Torrey Botanical Club, and stating that the enterprise had been announced and dis-
cussed at our meeting, and suggesting that the Club send a notice concerning this with
the field schedule to be issued in March, provided this date is not too late.

The scientific program consisted of two talks, the first by Dr. H. W. Rickett on
“The Genus Cornus in North America.”

The genus Cornus may readily be divided into 7 sections, 5 of which have oiten
been treated as genera. The difference between these are chiefly in the inflorescences.

CMV TE SOL Ee Cl UB 79

It is assumed that Afrocrania, with one species in East Africa is primitive. Closely
related is the big section Thelycrania, which covers much of Europe, Asia, and
North America, and is here typified by such species as C. stolonifera and C, amo-
mum. Also from Afrocrania came Tanycrania (C. mas, C. sessilis), found now in
southern Europe, China, western North America; Disocrania, with one species in
Mexico; Cynoxylon and Cephalocrania, which include such species as C. florida and
C. kousa, found in southern Asia and North America; and Arctocrania, the so-called
herbaceous boreal species C. canadensis and C. suecica. The progression seems to
have been from a primitive panicle subtended by bracts, by condensation to a “head”
with either disappearance of the bracts (Thelycrania), or their development into
more or less petaloid appendages; this often accompanied by the postponement of
anthesis through a dormant period until the season following flower-formation, the
bracts serving as bud scales. Most of the confusion in names and identities is in
Thelycramia. This section falls readily into groups of two or three species each, in
North America. A study of their distribution indicates that each of these groups
seems to have once been present in the southern Appalachian region, and to have
split as it migrated northward. When the segregated elements came again into con-
tact we find integrading forms which cannot be accurately classified. One of the
regions where this occurs is the Ohio Valley, where Rafinesque created numerous
new “species.” Another is the St. Lawrence Valley and northern New York.

This was followed by a presentation by Mr. F. R. Swift on “Treating Yeast Plants as
Individuals,” illustrated with splendid motion pictures.

This talk gave a short review of some of the methods used in developing yeast
cultures, from the primitive method of merely exposing easily fermentable material
to the air to the manipulator method developed at the Fleischmann Yeast Laboratory.

In the latter, glass cover-slips are pre-coated with a vegetable-mineral oil mix-
ture, adjusted to fit the medium in use at the time. Small hanging drops are then
distributed on the cover-slips and each one is seeded with one yeast cell. It was ex-
plained that by varying the proportions of the vegetable and mineral oil with the
varying surface tension of different media being used, easily handled, uniform drop-
lets, can be assured.

The development of yeast cultures growing and sporulating in such droplets was
shown in a series of slides and by stopmotion photography, in a motion picture.

The discussion of these papers was continued after the meeting was formally adjourned
at 5:05 p.m., while tea was generously provided by The New York Botanical Garden.

Epwin B. Matzke, CoRRESPONDING SECRETARY.

FEBRUARY 2. MEETING IN THE Museum or NaturAL Hisrory.

The meeting was called to order by the President, Dr. Robbins, at 8:15 p.m. At-
tendance 43. The minutes of the preceding meeting were accepted. Five new members
were elected to Annual membership. Dr. Seaver reported that the Auditing Committee
had found the Treasurer’s books in excellent condition. The report was accepted. Pres-
ident Robbins then read the names of those appointed to the various standing committees
of the Club. The scientific program was presented by Mr. G. L. Wittrock of The New
York Botanical Garden who spoke on “Local Plants Used by the American Indians,”
and illustrated these with colored slides. Aiter a discussion period the meeting ad-
journed at 9:40 p.m.

Honor M. HoitiincHurst, RECORDING SECRETARY.

FEBRUARY 17. MEETING at THE NEw York BoraNnicaL GARDEN.

The meeting was called to order at 3:30 p.m. by the President, Dr. Robbins. At-
tendance 22. The minutes of the preceding meeting were approved. Two new Annual
members and one Associate member were elected. The first speaker on the scientific

AVOERARGES Vs 7X

program was Dr. Frances E. Wynne who spoke on “Variability and Distribution of
Drepanocladus in North America.” 7 i

Drepanocladus, like many aquatic and semi-aquatic plants, is extremely variable.
Field and herbarium studies have been made to determine which variations are
hereditary and which are merely environmental fluctuations. Careful examination of
leaves from different parts of the same plant shows that elongated leaves, costae,
and cells are always produced when the plant grows submerged in water, whereas
shorter leaves and cells are produced by stems which grow emergent. Many of the
described varieties are merely seasonal phases produced by changes in the water
level. The present monographic study has reduced the previously recognized 24 spe-
cies and 30 varieties to 9 species, 1 subspecies, and 4 varieties. Hereditary factors
determine the presence of an excurrent costa and secund leaves; therefore these
characters are used as the basis for varieties in several species. The environmental
fluctuations of the shape of the leaves, costae, and cells are not given taxonomic
recognition.

The species of Drepanocladus may be classified geographically into two groups—
arctic-alpine and boreal-montane. The arctic-alpine species are restricted in their
range to the arctic regions; the boreal-montane species are widespread in the arctic
but occur also in boreal and mountain bogs and swamps.

The species of Drepanocladus may be divided into two categories on the basis
of fundamental variability. All the boreal-montane are extremely adaptable anc
variable and as a result of their toleration of a large variety of habitats have spread
over a wide range. The arctic-alpine species are stable, clear-cut species limited to
one region and one type of habitat.

Drepanocladus has a circumpolar distribution in both hemispheres. In North
America its present range coincides with the maximum extent of continental and
cordilleran glaciation during the Pleistocene. In eastern North America the distribu-
tion in partially glaciated states such as Pennsylvania, New Jersey, Ohio, Indiana,
and Missouri is significant. In these states Drepanocladus does not occur south of
the till sheets except in a few isolated stations. In western North America it is found
in mountain bogs and alpine meadows.

Four types of localities may have provided refuges for plants such as Drepanoc-
ladus during the Pleistocene: (1) areas south of the Pleistocene ice (2) arctic areas
north of the ice (3) unglaciated lowlands and (4) mountains or nunataks.

Two types of distribution result from the Pleistocene glaciation: (1) relic, static
and (2) general, widespread. Any hypothesis, proposed to explain the post-Pleisto-
cene dispersal of plants, must consider these two types of distribution found on
glaciated areas. Of the numerous explanations which have been proposed, the most
satisfactory is founded on the genetic constitution of the plants. Species may be
plastic and adaptable or rigid and static. The boreal-montane species of Drepanoc-
ladus are adaptable because a large number of individuals survived the Pleistocene
in a large variety of habitats‘on all of the possible refuges; therefore a large number
of biotypes contributed to these plastic species. The arctic-alpine species are rigid
because only a few individuals survived in a few habitats on only one of the refuges;
the biotypes contributing to these species were depleted by the vicissitudes of the
ice age leaving the species genetically rigid.

The second speaker was Dr. Morris Winokur who spoke on “Photosynthesis in Bac-
tenia.c

The attempt to interpret the metabolism of the green and sulfur bacteria has
resulted in the development of a generalized concept of photosynthesis which is
applicable to the green plant as well.

At the beginning of the twentieth century, there existed three conflicting theories
concerning the physiology of the purple bacteria. Engelmann believed that the purple
bacteria were able to photosynthesize much in the manner of the algae. Winogradsky
postulated that the oxidation of hydrogen sulfide and sulfur represented a substitute
for the respiration of organic substances, characteristic of the normal functioning of
most organisms. Molisch developed the thesis that the purple bacteria cannot assimi-
late carbon dioxide, but they assimilate organic compounds in the light. The contro-
versial nature of the results obtained by these three investigators was due to their

ACHIV Tipit sS yO kittie. CLUB 81

use of different biological materials: Engelmann—purple sulfur bacteria; Molisch—
purple non-sulfur bacteria; Wainogradsky—colorless sulfur bacteria. Buder at-
tempted to harmonize the diverse views by categorizing the organisms employed.
The existence of an intimate connection between the photosynthetic activity of the
purple sulfur bacteria and their respiratory phenomena was first clearly expressed
by Kluyver and Donker.

Van Niel demonstrated conclusively the photosynthetic nature of the metabolism
of the purple sulfur bacteria by devising methods for growing them in pure culture
in strictly mineral media in the light. His data show that the photosynthetic carbon
dioxide utilization depends quantitatively on the oxidation of sulfide and sulfur. He
also disclosed a similar relationship for the green sulfur bacteria. Comparing these
photosyntheses with that of the green plant, van Neil formulated the hypothesis
that the several photosynthetic reactions are all examples of photochemical carbon
dioxide reduction with a different hydrogen donor in each case. This generalized
view of photosynthesis made possible the explanation of the photosynthesis of the
non-sulfur purple bacteria as one in which the normal inorganic hydrogen donors
for the reduction of carbon dioxide are replaced by organic molecules. A variety of
indirect and direct experimental evidence has substantiated this interpretation.

Critical evaluation of the objections to the generalized concept of photosynthesis
leaves unimpaired the viewpoint that photosynthesis is a photochemical carbon
dioxide reduction in which organic compounds as well as inorganic substances or
even molecular hydrogen can play the role of hydrogen donors.

The consequence of the acceptance of this broad generalization is that it renders
untenable the classical Willstatter-Stoll theory of green plant photosynthesis.

After discussion of both papers, the meeting adjourned at 4:35 p.m., to be continued
over the inviting tea and refreshments served by friends at the Garden.

Honor M. HoLitinGHuRST, RECORDING SECRETARY.
Marcu 2. MEETING IN SCHERMERHORN EXTENSION, COLUMBIA UNIVERSITY.

The meeting was called to order at 8:15 p.m. by Dr. Lela V. Barton, the second
Vice-President. Despite the promise of a five siren air raid drill, 17 members attended.
The minutes of the preceding meeting were approved. The scientific program was pre-
sented by Dr. Ray F. Dawson who spoke on “Some Aspects of Parasitism in the
Mycorrhizae of Shortleaf Pine.”

The fungus or fungi which induce mycorrhiza formation on the roots of shortleaf
pine in the Missouri Ozarks area are apparently obligate parasites. The nature of
the symbiotic relationship between fungus and tree roots is determined largely by
environmental factors. When the trees are grown upon soils which are nutritionally
poor or unbalanced or when light intensity is low, fungal invasion of the short roots
readily occurs, and many well developed mycorrhizae are formed. Tree growth may
vary from slow to negligible. When the trees are grown upon fertile soil mycorrhiza
formation is difficult and slow and tree growth may be good, but if the soil contains
appreciable amounts of organic matter the seedlings will most likey fall victim of
damping-off fungi. When the trees are grown upon soils which contain relatively
low amounts of the necessary nutrients but when these nutrients are present in
physiologically balanced proportions on the soil colloids mycorrhizal development
and tree growth are both favored. Under such circumstances the mechanism of the
beneficial effect of mycorrhizae upon tree growth seems to be associated with an
increased salt absorption which is conditioned by an increased rate of aerobic respira-
tion and by a newly introduced mechanism for anaerobic respiration both of which
serve to maintain the energy output necessary for the growth processes. Hydrogen
ion excretion by the roots under such circumstances is increased several times thus
making it possible for the root colloids to undergo more intensively base exchange
reactions with the soil colloids in the initial phase of salt absorption. The enhanced
absorption of salts may then bring about greater water absorption and resulting in-
creases in both volume and mass of the plant tissues.

Following the discussion period, the meeting adjourned at 9 :35 p.m.

Honor M. HoiitincHurst, RECORDING SECRETARY.

MOR RE YA

Marcu 17. MEeetinc at THe New York BoranicaLt GARDEN.

The meeting was called to order by the Vice-President, Dr. Seaver, at 3:30 p.m.

Attendance 36. The minutes of the preceding meeting were accepted. Dr. Whaley an-
nounced the death of Dr. Tracy Hazen on March 16th, in Waterbury, Conn. On a
motion by Dr. Stewart it was voted that the Secretary send the condolences of the Club
to the family of Dr. Hazen. Dr. Matzke stated that he had received from Dr. Moulton,
of the American Association for the Advancement of Science, a request for a summary
of the history of the Torrey Botanical Club. Dr. Matzke said he would be willing to
prepare this summary which is to be published in the Journal of the Association with
the histories of other affiliated societies. The first scientific paper was presented by Dr.
Ernest Naylor who spoke on “Problems of Cellular Behavior during Regeneration.”

The author presents a brief discussion of some of the cell changes during early
stages of shoot and root formation on isolated plant parts during regeneration. The
multiplication and organization of cells during regeneration involves two funda-
mental types of cells morphologically. One is the meristematic type, which may or
may not be definitely organized into recognizable growing points. Such cells may be
variously located in leaf axils, nodai regions, leaf margins, woody structures, and
in other places.

The other type is concerned with differentiated cells of the plant body which
undergo structural changes and become actively meristematic to produce the new
root and shoot primordia. Such de-differentiation of vacuolate cells is described in
various tissues of a number of seed plants. The extent and limitations of such de-
differentiations in plant cells is briefly considered and some of the theoretical im-
plications pointed out.

Dr. Whaley was the second speaker on the scientific program, and his topic was “In-
feriority Complexes in Plants.”

Recent work of Dobzhansky and others indicates that in natural populations
many detrimental recessive genes are accumulated. The number and relative potency
of these genes is dependent upon the population structure, which is a function of the
number of individuals and the type of reproductive mechanism. Under selection it is
also possible for unfavorable dominants to accumulate. Heterosis is the result oi
masking of these deleterious recessives in some organisms, the result of heterozy-
gosity in others. Suggestions as to the nature of some of. these deleterious factors
is found in excised root culture experiments. The roots of certain tomato lines show
a deficiency in ability to synthesize pyridoxine, others in the ability to synthesize
nicotinamide. Crosses between such lines produced vigorous hybrids under ordinary
field conditions. Hybrid vigor represents a return to an “optimum” phenotype rather
than any “super” phenotype.

After a discussion of both papers, the meeting adjourned at 4:35 p.m. Then tea and
delicious refreshments, in keeping with the spirit of St. Patrick’s Day, were served by
friends at the Garden.

Honor M. HoLiincHurst, REcoRDING SECRETARY.

Marcu 27. Fietp Trip to The New York Zoological Park for the study of some animal

habits. Leader, Miss Nellie L. Condon, Director, Reptile Study Society of America.
Attendance 11.

Marcu 28. Fretp Trip to Springdale, N. J., for limestone lichens. Leader, Mr. G. G.

Nearing. Attendance 4. Unusual forms found were: Acarospora murorum, Cypheliun
tigillare, and Physcia venusta. The last two were in fruit, and these fruiting forms
appear to be rare.

Aprit 4. Fretp Trip to Central Park, N. Y. to search for the trees mentioned in L. H.

Peet’s book “Trees and Shrubs of Central Park’ (1903). Leader, Dr. E. B. Matzke,
Columbia University. Attendance 25. Many of those present took an active part in the

FNC APIUNAMIPIUES) (OT Wiest, CIOs} 83

identification of the trees, and all of us profited by Mr. James Murphy’s generous and
genial contributions on the trees and shrubs as well as on the history and lore of Central
Park. The following plants were found in flower: Cornus mas, Lonicera fragrantis-
sima, Ulmus americana, U. campestris, and Acer rubrum. Most of the day was spent
trying to locate trees mapped in Peet’s book. The morning was devoted to the southern
half of the park, and the afternoon to the northern end. A few tentative conclusions may
be suggested, subject of course tc correction after more careful study:

1. Changes in tree population have been much more pronounced in the southern
end of the Park in the last forty years than in the northern end. Many of the trees
listed by Peet for the northern end could easily be located; this was decidedly not
true nearer 59th Street.

2. The conifers have not fared well. White pines and some other gymnosperms
present in 1903 were not found; a young Douglas Fir, more recently planted, was
distinctly the worse for wear. Some Austrian pines have survived, and they may or
may not be an exception.

3. The Turkey Oak, Quercus Cerris, has grown and perhaps prospered; native
oaks apparently do not thrive.

4. The English Elm, Ulimus campestris, seems to have done reasonably well,
distinctly better than our native ones.

5. In the wetter habitats the red maple, Acer rubrum, seems tq be pretty well
established.

6. Ailanthus apparently “seeds in” in the park.

Epwin B. Matzke

APRIL 6. MEETING IN SCHERMERHORN HALL, CoLtumMBIA UNIVERSITY.

The meeting was called to order at 8:20 p.m. by the President, Dr. Robbins. At-
tendance 33. The minutes of the preceding meeting were approved. Eight new annual
members and three associate members were elected, and one transfer from associate to
annual membership was approved.

Dr. Matzke then read to the club the letter which he, as Corresponding Secretary,
had sent to the family of the late Dr. Tracy E. Hazen:

New York City
March 23, 1943
Dr. Ropert HAZEN
Thomaston
Connecticut.

Dear Dr. Hazen:

At its meeting held on March 17, 1943, the Torrey Botanical Club directed its
Secretary to extend sympathy and condolence to the family of the late Professor
Tracy Elliot Hazen.

Its Editor for many years, its President for two terms, the Torrey Botanical
Club was singularly fortunate in having profited by the sound scholarship, the
meticulous labors, the faithful devotion to duty, and the kindness of heart of Pro-
fessor Hazen. All its members admired him, all respected him as a thorough gentle-
man, and all who knew him intimately, loved him.

Your grief, and ours, may be assuaged by a knowledge of Professor Hazen’s
goodness, of his quiet nobility, and of his high attainments.

The Torrey Botanical Club realizes that a faithful officer, member, and friend
has passed to his reward; it is grateful for having shared in the innate richness of
his life.

In deep respect,
Epwin B. Matzke,
CORRESPONDING SECRETARY.

84 TORREYA

Dr. Robbins announced that he had appointed a committee to draft a biographical note
on Dr. Hazen for the BULLETIN. The committee consists of Dr. Carey, chairman, Dr.
Barnhart, and Dr. Bold.

The scientific program oi the evening was presented by Dr. A. B. Stout of The
New York Botanical Garden, who spoke on “Dichogamy in Relation to Reproduction,”
illustrated with lantern slides. After questions and discussion from the floor, the meet-
ing adjourned at 9:40 p.m.

Honor M. HoLitincHurst, RECORDING SECRETARY.

Aprit 11. Fretp Trip to The Brooklyn Botanic Garden and Conservatories for seasonal
studies outside, and for observation of economic plants from other lands. Leaders,.
Dr. A. H. Graves and Dr. A. Gundersen of the Garden staff. Attendance: 3 members
and 147 visitors in the Garden.

Aprit 17, Fretp Trip to The New York Botanical Garden Conservatories, particularly
the Easter exhibit in the Display House featuring trees of the Holy Land. Leader, Dr.
H. A. Gleason of the Garden staff. Attendance 11.

Aprit 18. Fretp Trip to the Lichen Trail in Palisades Interstate Park for lichens, fungi,
and general botany of the season. Leader, Mr. G. G. Nearing. Attendance 4.

Aprit 21. MEETING aT THE BROOKLYN Botanic GARDEN.

The meeting was called to order at 3:30 p.m. by Dr. C. Stuart Gager, in the absence
of the President and Vice-Presidents of the Club. Attendance 18. The minutes of the
preceding meeting were approved. The program consisted of a talk by Dr. Henry K.~
Svenson on the “Plants of a Long Island Pond,” illustrated with Kodachrome slides ;
and an inspection of the Local Flora Area of the Garden under the leadership of Dr-
Svenson.

Honor M. HoLtincHurRST, RECORDING SECRETARY.

Aprit 24. Fietp Trip to Surprise Lake, Watchung Reservation, near Summit, N. J., for
reptiles, amphibia, and spring plant life, all of which reflected the late season. Leader,
Miss Nellie L. Condon. Attendance 11.

May 1. Fretp Trip to Mertensia Island along Raritan River above Raritan, N. J., to see
the profuse stand of Mertensia, Dentaria, Erythronium, etc. This was the ideal date for
this season. Leader, Dr. John A. Small, New Jersey College for Women. Attendance 6.

May 2. Fietp Trip to Silver Lake, White Plains, N. Y., for spring flowers and birds.
The day was cold and windy: 25 bird species were seen, 4 violets and 10 other plant
species were found in bloom. Leader, Miss Farida A. Wiley, American Museum of
Natural History. Attendance 12.

May 8-9. WEEK END FIELD Trip to Camp Thendara, Lake Tiorati, Palisades Interstate
Park, N. Y., for study of birds and plants. Leader, Mrs. Richard M. Abbott. Attendance
32, of which at least 5 were from the Torrey Club. 69 bird speceis were recorded,
including 19 warblers.

May 11. MretiInc in SCHERMERHORN Hatt, CoLuMBIA UNIVERSITY.

The meeting was called to order by the President, Dr. Robbins, at 8:15 p.m. At-
tendance 70. The minutes of the preceding meeting were approved. The scientific pro-
gram was presented by Dr. Samuel Record, of the Yale School of Forestry, who told
“How Woods Are Identified.”

The results already obtained from the systematic studies of woods indicate

clearly that any wood sample is identifiable. The unit of classification is at present
the genus, but well-defined species are frequently recognizable and their number

IMG IMIWAN I WS) (Ole IMs tes, (CHAVIS) 85

The essentials for systematic study of woods are: 1. A comprehensive and repre-
sentative collection of samples obtained with herbarium material determined by
competent taxonomists. In the Yale collections there are 40,700 catalogued samples
representing nearly 12,000 named species of 2,800 genera and 232 families. 2. A col-
lection of slides with cross, radial, and tangential sections for examination under the
microscope. The Yale slide collection contains about 19,500 slides of 11,072 specimens
of 6,506 name species, 2,616 genera, and 218 families. 3. Careful examination of the”
slides by trained anatomists and the preparation of descriptions and tabulations of
all essential features. The standards used are those approved by the [International
Association of Wood Anatomists after several years of cooperative effort. 4. The
use of the assembled data for making keys or other aids to identification. Numerous
keys to special groups have already been published and others are in preparation.

The task is very large, difficult, and costly and can only be carried out success-
fully through cooperative efforts. Ordinary taxonomists, though willing to accept
the aid of the anatomist in a time of trouble, make no effort to secure material
essential for anatomical study. Fortunately there are exceptions to this rule and
The New York Botanical Garden is foremost among American institutions in en-
couraging its botanists to collect wood samples. Systematic wood anatomy has made
its greatest progress during the past decade for the simple reason that during that
time research workers in various parts of the world effected an organization and
pooled their efforts and materials. The best incentive to further progress would be
the addition of new and better material which botanical expeditions could so readily
supply.

Because discussion of the talk was sharply curtailed at. 9:30 p.m. by the sounding of
sirens for an air raid drill, the meeting quickly adjourned to darkened halls, where by
the light of a lantern, Dr. Record graciously identified wood specimens presented by
members of the audience.

Honor M. HoLLtinGHuRST, RECORDING SECRETARY.

May 15. Fietp Trip to McLean Woods, The Bronx, N. Y., for spring study of the area.
Species lists were prepared and filed with the Field Committee. Leader, Mrs. Mary
Holtzoff. Attendance 14.

May 16. Fretp Trip to Point Pleasant, N. J., to search for Britton’s Violet, of which a
good stand was found, and in addition a large number of other plants. Leader, Mr. Louis
Hand. Attendance 7.

May 21-23. Fietp Trip to Culvers Lake, N. J., for the Annual Branchville Nature Con-
ference. In order to have the conference at the most desirable season and without in-
creased expense to those participating it was necessary to change from THE PINES
to THE HALTERE for accommodations. This proved satisfactory. Leaders: Mr.
Wallace M. Husk, Professor Oliver P. Medsger, and Dr. Julius Johnson. Attendance
40.

May 22. Fretp Trip to Ridgewood, N. J., to see the Rhododendron seedbeds, nurseries, and
stock of the leader, Mr. G. G. Nearing. Attendance 10.

May 29. Fretp Trip to Haskell, N. J., for fungi of the Chicohikie Falls region, especially
Fissipes acaulis, of which there was plenty. Leader, Mr. F. R. Lewis. Attendance 5.

aN t 5

A Past

Vor. 43 APO) IR IRI, WE aN Jury 1943

The Importance of Taxonomic Studies of the Fungi*

Frank D. Kern

The naming and classifying of living organisms has been going on for
centuries. It has been well said that “a large part of our thinking about living
things is bound up with some system of classification.” Another writer has ,
pointed out the fact that we depend much upon classification in our general
experiences. “It is the innate propensity of active minds,” he says, “to form
species, 7.e., successively to make distinctions, to point out similarities, and then
to assemble the things that are alike into their kinds. It applies to everything
from chemical elements to college fraternities.”

* Read at the 75th Anniversary Celebration of the Torrey Botanical Club at The New
York Botanical Garden, Tuesday, June 23, 1942. Contribution from the Department ot
Botany, The Pennsylvania State College, No. 137. Publication authorized on July 6, 1943

as paper No. 1185 in the Journal Series of the Pennsylvania Agricultural Experiment
Station.

66 TORRE VEX.

The early workers who studied the microfungi under the microscope rather
naturally tried to interpret them in the light of their knowledge of the parts of
flowering plants. In the case of the bread-molds the sporangia seemed like
little fruiting pods containing seeds. By analogy rust spores were similarly
interpreted although the situation there was not so easily demonstrated as with
the molds. In 1807 DeCandolle, referring to the spores of Uromyces and
Uredo, said that “with a microscope this powder seems composed of ovoid or
globular spores .... filled with many small grains that are considered spores.”
He thought that a teliospore might contain at least 100 such “spores.” This
interpretation prevailed among such workers as Fries, Léveillé, and the
Tulasne brothers, and persisted until the time of De Bary in the middle of the
19th century.

The first author to make a distinct advance in the classification of the fungi
after the beginning of binomial nomenclature was Persoon. In a paper
published in 1794 (Neuer Versuch einer Sytematischen Eintheilung der
Schwamme, Romer’s Neues Mag. Bot. 1: 63-128) he recognized 77 genera
of fungi, which he placed in two classes: Angiothecium and Gymnothecium.
The three genera of rusts, which were included, were the first rust genera to be
established after the solitary rust genus of Micheli 65 years before. Several
authors of important works during the first quarter of the nineteenth century
followed Persoon’s classification in the main. Among these were Schumacher,
Rebentish, Albertini and Schweinitz, De Candolle, and Brongniart. During
the same period Link brought out a new classification which was accepted
- wholly or in part by Schlechtendal, S. F. Gray, and Wallroth.

KERN: TAXONOMY OF THE FUNGI 67

During the middle of the nineteenth century great contributions to the
knowledge of the larger fungi were made by Elias Fries. He had “not only a
_poor opinion of the parasitic fungi but an antiquated conception of their
nature.” In his third volume of “Systema Mycologicum’” (1832) he used the
name Hypodermii to include the rusts, smuts, and some other fungi and
characterized them as having “No proper vegetative body ; sporidia originating
from the metamorphose of the cellular structure of living plants: an inferior
kind of fungi.” Nevertheless the work of Fries which extended over more than
a half a century gave a great impetus to the study of fungi. His prestige was so
great that there were many who accepted his leadership. Among these may be
mentioned Endlicher, Leveillé, Corda, Rabenhorst, Strauss, Berkeley, and
Cooke. Most of these authors made changes in the arrangement of the genera.
Corda’s extensive publication (Icones Fungorum) is notable not only for its
contribution to the knowledge of the structure of the larger fungi but also for its
advances regarding hundreds of the microfungi.

During the first three quarters of the nineteenth century new species
were being recognized and named from all parts of the world. The descrip-
tions appeared in journals, reports, and books many of which were not widely
circulated. It is little wonder that investigators soon found it difficult to know
whether or not a species under consideration was already described and
named. It may be well said that this condition still exists. Thus it came about
that species were named and renamed from several to many times. Little was
known of the distribution of the fungi and workers in one region had no way
of knowing of the probability of the existence elsewhere of the species which
they were studying. Conceptions of the probable cosmopolitan distribution of
the fungi were necessarily slow in developing. Many efforts were directed
toward bringing together all species known to occur in certain regions or
countries without attempts to determine their wider distribution. The flora-
type of publication became common, especially in the European countries.
Rabenhorst’s ““Kryptogamen Flora” of Germany, Austria, and Switzerland
is a good example. Many other floras could be cited. These publications were
valuable but they did not solve the problem for the workers who were located
away from the European centers of mycological activity.

The assertion that many mycologists actually were deterred “from describ-
ing supposedly new species for fear of duplication” will doubtless not meet with
credulity. An important step toward overcoming this situation was the plan
for the “Sylloge Fungorum” inaugurated by Saccardo in 1882. The first
volume appeared in that year. The effect was an immediate stimulation of sys-
tematic mycological activity. This great work developed into twenty-five
volumes, the last appearing in 1931. During this period mycological journals

68 DAO MRR a

were established in various countries and taxonomic work with the fungi went
forward at a rapid rate.

Thus far we have given consideration chiefly to the describing, naming and
classifying of the many and varied forms. The earlier workers naturally were
concerned with these phases of study. It should not be concluded, however,
that there were not some, even among the early workers, who were intrigued
with the possibilities of studying the development and life-histories of the
forms with which they worked. There were suggestions that relationships
might exist between different forms which were found in close association.
The impress left by De Bary on this phase of mycological work is well known.
He began his work about the middle of the nineteenth century and the type
of investigation which it stimulated has continued up to the present. He found
time to work not only with fungi but also with algae, myxomycetes, bacteria,
and higher plants. It is said that no less than 68 workers, afterwards distin-
guished in science, studied under him at Strassburg. According to Erwin F.
Smith, “His work and that of his students put plant pathology on a new
foundation, and he also, undoubtedly had much influence on human and ani-
mal pathology, since his very successful infection experiments with fungi on
plants suggested many things to those who were trying to determine the cause
of human and animal plagues.’ Yet we must agree that the primary interest
of De Bary was in morphology rather than in pathology.

Using a good microscope and employing micro-chemical reagents De Bary
made important advances in the knowledge of spores, infection, and mycelia.
His cultural demonstration of heteroecism in Puccinia gramuinis, with proof
that the aecidium on barberry was a stage in the life-cycle of wheat rust is well —
known. These results were announced in 1865. This work, and more which
followed, ushered in a new phase of mycological endeavor. It is significant that
he began these investigations not out of pure scientific interest, but in order
to settle controversies between agriculturists and botanists regarding the rela-
tion between smuts and rusts and diseases. Agriculturists thought them to be
the causes of disease while botanists were inclined to regard them as products
of disease. De Bary had himself resisted the suggestion of a possible alternation
of generations which required an alternation of hosts plants. When his experi-
ments led to that conclusion, his naive statement that “one comes around, per-
haps, in a way, to the ancient opinion according to which rusted wheat would
be infected by the rust of barberry” is most interesting. His experiences should
be heartening to many present-day investigators who are required to work on
projects which are economic and agricultural in nature. Out of such problems
may arise basic scientific discoveries as in the case of De Bary.

The next epoch in the study of the fungi after De Bary was ushered in by
the study of the nucleus and its behavior. This gave a new direction to the

KERN: TAXONOMY OF THE FUNGI 69

study of fungi. As life-histories were important for taxonomic considerations
so nuclear developments were eventually recognized as having a bearing on
taxonomy. The application of cytological methods to the study of life-histories
in the fungi began with the work of Dangeard in 1894 and was soon under
way on a large scale. Other early workers in this field were Poirault, Sappin-
Trouffy, Maire, Harper, Blackman, and Christman. It was soon evident that
the nature of sexual reproduction in the fungi was of great value in determin-
ing relationships. We are indebted to such a host of investigators that it is
impossible to mention them by name. Notable studies have been made in the
Phycomycetes, Ascomycetes, Ustilaginales, Uredinales, and higher Basidio-
mycetes. In the last few years genetical studies have been made and highly
important results are in the making.

Our account would not be complete if we did not make some reference to
the possibility that the classification of the future may have a physiological
basis. Much headway toward such a goal has been made by Mez and his asso-
ciates. Many of you are familiar with the fact that Mez, using serological
methods, has constructed a family tree of plants which corroborates in a
remarkable manner the older tree based on morphological characters. Seifriz
refers to this work in a recent book (The Physiology of Plants, 1938) with the
remark, “It is of great significance to the field of evolution and phylogenetic
relationship that a purely chemical basis of classification should so well sup-
port a purely anatomical one.’’ Seifriz points out that the relationships between
plants established thus far by serology hold well for families, not so well for
genera, and not at all for species. He believes, however, that this is due to a
lack of delicacy in technique. He is of the opinion species differences in proteins
must also exist.

Our historical sketch which began with the early attempts to classify fungi
led us rather inevitably to some consideration of morphological, cytological,
genetical, and physiological studies. Certainly we must agree that knowledge
gained in all these fields is essential for progress in taxonomy. E. A. Bessey in
1939 (A Textbook of Mycology) refers to the present-day activity of sys-
tematic mycologists and points out that, “Life histories are being studied in
all groups, the sexual relations are being scrutinized from the lowest to the
highest fungi and genetical studies are revealing results somewhat parallef,
but on a vastly smaller scale as yet, to those attained by the study of Zea mays
and Drosophila.” ‘As never before,’ says Bessey, “is a knowledge of fungi
themselves so necessary.” Obviously right conceptions of. fungi must be based
upon many facts, and wrong conceptions can easily be the result of partial facts,
and of ideas derived from other plants which may be inapplicable and mis-
leading.

'~ We have referred to the contribution which Darwin’s theory of evolution

70 TORREYA

made to biological classification. Phylegeny soon became the fundamental basis
for classificatory endeavor. So far as the fungi are concerned we should not
overlook the influence of the work of Hofmeister in 1851 on the bryophytes
and pteridophytes. The recognition of an alternation of generations in these
groups had its effect on studies of the algae and fungi.

Every student who has taken a course in general botany is familiar with
the system of classification which places the algae and fungi together in the
division Thallophyta. We have no thought of attempting to reach any con-
clusions about this broad question of the taxonomic disposition of the fung.
Whether the fungi are to be regarded as one of two subdivisions of the Thallo-
phyta, the algae being the other, depends upon the origin of the fungi. We say
this in spite of a recent assertion that the taxonomist “‘is not interested in the
origin, but in the character of his plants.” On the origin of the fungi, G. M.
Smith, in his “Cryptogamic Botany,” Vol. I, “Algae and Fungi’ (1938)
writes, “This is highly controversial and opinion is divided as to whether
they arose from the protozoa or whether they had either a monophyletic or
polyphyletic origin among the algae. If they arose from protozoa, they should
be put in one or more divisions coordinate in rank with the various algal
divisions ; if they arose irom the algae, they should be placed as classes of one
or more oi the algal divisions.”

Smith reviews the algal and the protozoan theories of the origin of the
fungi and concludes that “it seems more probable that the fungi evolved from
protozoa rather than irom algae.” He bases his conclusion largely on metabolism
and the type of flagellation in the Phycomycetes. There are some algal groups
in which there occur chlorophyll-less forms which are so similar morpho-
logically that they cannot be regarded as distinct from the green forms. It is
pointed out that these saprophytic and parasitic algae accumulate reserve
carbohydrates as starch just as do the green algae. In contrast the Phycomy-
cetes are reported generally to accumulate carbohydrates as glycogen but
never as starch. The zoospores and gametes oi the green algae are never uni-
flagellate whereas the motile cells of certain Phycomycetes are regularly uni-
flagellate. It is admitted that the question of the origin of the Ascomycetes is a
more difficult one. The similarity in the sex organs, and the structures developed
subsequent to fertilization, in the Ascomycetes and in the red algae are strik-
ing and have caused many workers to assume a relationship between these
groups. Smith argues that these distinctive reproductive structures may have
evolved along independent phyletic lines. He thinks the Ascomycetes had
their origin in the Phycomycetes and that the Basidiomycetes arose by modi-
fication from the Ascomycetes. In his classification he rejects the Thallophyta _
as a division of the plant kingdom and in its place substitutes nine divisions,
of which the Myxothallophyta, or slime molds, constitute one and the

KERN: TAXONOMY OF THE FUNGI 7

Bessey in his “Textbook of Mycology” has attempted a definition of the
term fungi that would not commit the definer to any system of classification.
We quote: “Fungi are chlorophyll-less thallophytic organisms typically con-
sisting of coenocytic or cellular filaments, but including also encysted or
amoeboid one-celled organisms which reproduce by some type of motile or
non-motile spore; excluding the Bacteria and such chlorophyll-less organ-
isms, which, by their structure, are with definiteness assignable to recognized
orders of algae.”” Bessey is of the opinion that the Mycetozoa are not related
to the fungi; are not, indeed, plants. There are those who believe that the fungi
should not be regarded as belonging to the Plant Kingdom. Herbert F.
Copeland in a comparatively recent paper (Quarterly Review of Biology,
December, 1938) has presented evidence and argument “to the effect that
organisms can be arranged, naturally, and more conveniently than in the
past, in four Kingdoms as follows”:

Kingdom 1. Monera (Bacteria and Blue-green Algae)
Kingdom 2. Protista (Protozoa, Diatoms, Red and Brown Algae, Slimemolds, and

Fung1)
Kingdom 3. Plantae (Green Algae, Liverworts and Mosses, Ferns and Allies, Seed
plants )

Kingdom. 4. Animalia (Metazoa)

To those who have been accustomed to thinking that all living organisms
must be either plants or animals the recognition of two new groups as King-
doms may seem revolutionary. It is true, however, that the line between lower
plants and lower animals has always been a difficult one to draw. It must
be admitted that nomenclatorially there are difficulties in placing together in
the Kingdom Protista organisms which have been previously in two different
Kingdoms. The original proposal for a Kingdom to be called Protista was made
by Haeckel in his ‘“Generelle Morphologie” in 1866. He also established the
group Monera but included it in Protista. According to Copeland other
authors have expressed the opinion that the Monera should be treated as a
separate Kingdom.

72 WO RARE YN

the fact that there is no general agreement as to whether the fungi are
monophyletic or polyphletic in origin or whether they have descended from
the algae or from the protozoa. The algal theory appears to have been advocated
by A. Braun in 1847, and was accepted by Cohn (1854), Pringsheim (1858),
and Sachs (1874). De Bary in 1881 objected to the method of intercalating
the fungi among the algae saying it led to an orderly arrangement of species
but not to a natural system. The suggestion that the fungi arose from the
protozoa is credited to Cornu (1872), and was developed by Gobi (1885)
and Dangeard (1886). Atkinson (1907) was in favor of deriving the lower
fungi from ancestral unicellular organisms, but was uncertain whether they
were colorless or chlorophyll bearing. He was, however, certain that their
origin was monophyletic. The algal origin of fungi was supported by Stras-
burger and C. E. Bessey. Gauman (1925) presented the view that all true
fungi were derived from the green algae in monophyletic line; he believes the
lower Chytridiales (his class Archimycetes) along with the Myxomycetes may
have arisen from the colorless Flagellatae. He does not regard either of these
groups as fungi. Martin (Bot. Gaz. 93: 421-435, 1932) has “suggested that the
fungi be regarded as a phylum which has not definitely developed into either
plants or animals, but may be grouped with the former as a matter of con-
venience, and in accordance with custom.” He rejects the assumption that
all living organisms are descended from a single primitive cell and points
out that the assumption that life may have originated more than once and
in different forms is more in accord with what we know of living organisms.

KERN: TAXONOMY OF THE FUNGI

a |
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As with other groups of living organisms the fungi have had _ their
devotees. Crowds of them have advanced to the expert stage. It is impossible
to name them or to evaluate their contributions. They must be treated
generically, as it were. The writer has thought it worth while to try to present
some of the problems which such workers encounter. By this is meant not so
much the problems inherent in taxonomic studies but rather the wider limita-
tions which often operate to check individual progress and to break the con-
tinuity of advances for which a groundwork may have been well established.
The difficulties which are to be discussed are not necessarily peculiar to sys-
tematic mycology. Taxonomic work in general as well as in mycology, has
a checkered history. Its advances through the centuries have been piecemeal.
Perhaps it will always be thus, and deploring the fact may not only be in
vain but may not be fitting.

Even where universities, colleges, or other institutions or governmental
agencies are involved it is still true that the ambition, industry, and perseverance
of individuals are largely responsible for the advances that have been made. In
these later days we have been hearing a good deal about institutional research.
So far as taxonomic work with the fungi is concerned we delieve that an
analysis would show that research in this line is mostly due to individual
prosecution rather than to institutional initiation. It may, happen that an
institution will make an effort to continue the type of research that has been
inaugurated and successfully carried on by one of its staff members and will
then refer to the program as an institutional program. More often it happens
that a real leader appears and develops successfully a line of work which 1s
supported (more or less) during his years of activity but which is dropped
by the institution afterwards. Such instances indicate the correctness of the
conclusion that there is often no such thing as an institutional program. There
are, of course, exceptions but we feel safe in saying that the exceptions prove
the rule rather than make it. We have inserted the parenthetical phrase—
more or less—because we are sure that institutional support even when

74 AMOURGREE YSN

Someone may well ask why these difficulties are raised in connection
with taxonomic research when they exist in so many lines of research activity.
There are several reasons for doing so. The source materials for taxonomic
research are in large part not commercial commodities. They consist of rare
books, separates, indexes, illustrations and specimens which are accumulated
only with time, patience, correspondence, and exploration. When such col-
lections have finally been put together in an institution they should be used
by more than one generation of workers in that institution. Or if that is not
possible some method should be worked out by which they become available
to succeeding investigators in other institutions. There are now in existence
some collections of microfungi where spore measurements and drawings
accompany literally hundreds of specimens. Such aids are indispensable for
taxonomic studies and when available not only save the time necessary to
duplicate them elsewhere but help to prevent errors and misconceptions.
There are also herbaria of fleshy fungi where great accumulations of photo-
graphs, drawings, and notes make them of the utmost importance to other
workers. This is not a plea for the centralization of mycological taxonomy.
It is rather to call attention to the fact that enormous resources are fre-
quently accumulated and then not used nor made available for use. Since our
modern concepts fix the application of names by types rather than by descrip-
tions it is a fair question whether type specimens should ever be personal
or institutional property. The difficulties may seem insurmountable but this
may not be the case. Surely we will make no progress until the workers them-
selves reach a keener appreciation of the situation.

KERN: TAXONOMY OF THE FUNGI i 75

A few weeks ago I received a letter from a former associate in which he
said, “I notice, with much interest, in the last issue of Science, that you are to
have a part in the “Symposium on Taxonomy,’ June 23, in connection with the
Seventy-fifth Anniversary Celebration of the Torrey Botanical Club... . I
assume that you will speak for the fungi.” Of course. Whether I have said, or
still can say, anything which he would have me say is another matter. I assume
that he expected me to make some reference to the problem of nomenclature
and it seems impossible to close this discussion without bringing up this
vexatious topic.

I propose to make comments of a general nature and to confine them to

- two aspects of the nomenclatorial situation: (1) on getting rules, and (2)
on getting them into effect.

76 TORREYA

fundamental principles of a basic nature in which they believed were not -

incorporated. They felt that once they accepted a code without these principles
the chances for amendment would not be good. I have in mind chiefly the
“type-concept” which was not a part of the original code. Reference to a more
or less minor feature may serve to illustrate difficulties regarding adoption.
The Vienna code provided that “On and aiter January 1, 1908, the publica-
tion of names of new groups of recent plants will be valid only when they are
accompanied by a Latin diagnosis.” Again I do not know how many names
have since been published which are invalid, but I do recall taking part in a
business session of a certain mycological society, at least 25 years after the
' Latin deadline, when the matter before the house was whether that rule
should be enforced in its official journal.

It seems fair to say that cordial agreement was reached at the Cambridge
Congress in 1930 on most of the disputed nomenclatorial problems and that

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KERN: TAXONOMY OF THE FUNGI 77

the disposition to accept International rules was improved thereafter. Not long
ago I was criticized by a colleague for such a conservative statement. He wanted
me to say that these rules are, and have been for some time, actually in effect.
Again it may be time which settles many problems. At any rate, it was in
1940 that the Secretary of the United States Department of Agriculture
formally approved a recommendation of the Department Committee on Plant
Names “to put the Department, botanically speaking, under the International
Rules of Nomenclature.” To me it is interesting that it took ten years for
this department to come to an action making these rules official for “publica-
tions, reports, and correspondence involving scientific plant names.’ Perhaps
one might be pardoned for calling attention to the anomaly of an agency finally
finding it expedient to subscribe to the acts of an organization which it failed
officially to aid. It is also interesting to note that two years after the official
order they are still going through an adjustment period in getting nomen-
clatorial usage realigned according to International rules. When it becomes
necessary to drop the name Ustilago hordei which, according to old usage,
has been applied to the covered smut of barley and to take up the same name,
according to International Rules, for Joose smut of the same host it 1s little
wonder that the workers talk about confusion. Personally, I believe that the
confusion will be only temporary and that the advantage of getting on a
world-usage basis will more than outweigh the disadvantages. It is desirable
to avoid changes in names as far as possible, but changes cannot be entirely
avoided if the rules of nomenclature are to put in order the old names as
well as to be a guide for the creation of new names. There are those who
believe that the procedure embodied in the present system of nomenclature
leaves too much to expediency and personal preference and do not rest
sufficiently upon foundamental principles. It has been pointed out that “there
is no guarantee—if, indeed, there is any hope—that the system which may. be
adopted today will be accepted by the next generation.’’ No, there is no
guarantee that anything man devises will continue—not even democracy. We
must hot, however, look upon this or any other problem in such a futile
manner. There are difficulties, to be sure, but they are not insurmountable.
We are told in the Torrey Botanical Club Announcement and Field Schedule
for 1942, “It is understood that there will be no mutilation of species at this
session.” That being the case, this seems to be the proper place to bring this
discussion to an end.

THe PENNSYLVANIA STATE COLLEGE
STATE COLLEGE, PENNSYLVANIA

Voi. 43 ID QURIRAD VEEN Jury 1943

ACTIVITIES OF EEE CLUE

January TO May 1943

January 5. ANNUAL MEETING.

The annual dinner meeting of the Torrey Botanical Club was held at the Men’s
Faculty Club, Columbia University, at 6:45 p.m. The President, Dr. C. Stuart Gager,
presided, with 82 members and friends present. After the dinner the minutes of the
preceding meeting were approved. The reports of the Treasurer, of the Chairman
oi the Field Committee, and of the Editor of Torreya were distributed in mimeo-
graphed form, and the combined report of the Editor and the Bibliographer was read
by Dr. Matzke. These reports were accepted on a motion by Dr. Karling.

Dr. Gager addressed a few remarks to the Club and then announced that the following
list of officers had been elected for the year 1943:

President: William J. Robbins

Ist Vice-President: Fred J. Seaver

2nd Vice-President: Lela V. Barton

Corresponding Secretary: Edwin B. Matzke

Recording Secretary: Honor M. Hollinghurst

Treasurer: W. Gordon Whaley

Editor: Harold W. Rickett

Bibliographer : Lazella Schwarten

Business Manager: Michael Levine

Members of the Council: Charles A. Berger, Clyde Chandler, Albert E. Hitchcock,

Roger P. Wodehouse
Delegate to the Council of the N. Y. Academy of Sciences: Bernard O. Dodge
Representative on the Board of Managers of the N. Y. Botanical Garden: Henry
A. Gleason

Representatives on the Council of the American Association for the Advancement

of Science: John H. Barnhart, Albert F. Blakeslee

Honor M. HoLitincHurst, RECORDING SECRETARY.

January 20. MEETING aT THE NEW York BoTANICAL GARDEN.

The meeting was called to order at 3:30 p.m. by the President, Dr. William J.
Robbins. Attendance 25. The minutes of the preceding meeting were approved. The
iollowing new members were elected: 15 to Annual membership, 3 to Associate mem-
bership; 2 transiers to Annual membership and 4 transfers to Associate membership
were approved. The resignations of 21 Annual and of 4 Associate members were
accepted with regret.

A letter was read concerning the preservation of High Tor. Dr. Robbins suggested
that a letter be sent to the sponsors oi this movement, expressing the interest oi the
Torrey Botanical Club, and stating that the enterprise had been announced and dis-
cussed at our meeting, and suggesting that the Club send a notice concerning this with
the field schedule to be issued in March, provided this date is not too late.

The scientific program consisted of two talks, the first by Dr. H. W. Rickett on
“The Genus Cornus in North America.”

The genus Cornus may readily be divided into 7 sections, 5 of which have oiten
been treated as genera. The difference between these are chiefly in the inflorescences.

IME AMIDA S) Okt Was, (Cie Oi} 79

It is assumed that Afrocrania, with one species in East Africa is primitive. Closely
related is the big section Thelycrania, which covers much of Europe, Asia, and
North America, and is here typified by such species as C. stolonifera and C. amo-
mum. Also from Afrocrama came Tanycrania (C. mas, C. sessilis), found now in
southern Europe, China, western North America; Disocrania, with one species in
Mexico; Cynoxylon and Cephalocrania, which include such species as C. florida and
C. kousa, found in southern Asia and North America; and Arctocrania, the so-called
herbaceous boreal species C. canadensis and C. swecica. The progression seems to
have been from a primitive panicle subtended by bracts, by condensation to a “head”
with either disappearance of the bracts (Thelycrania), or their development into
more or less petaloid appendages; this often accompanied by the postponement of
anthesis through a dormant period until the season following flower-formation, the
bracts serving as bud scales. Most of the confusion in names and identities is in
Thelycrania. This section falls readily into groups of two or three species each, in
North America. A study of their distribution indicates that each of these groups
seems to have once been present in the southern Appalachian region, and to have
split as it migrated northward. When the segregated elements came again into con-
tact we find integrading forms which cannot be accurately classified. One of the
regions where this occurs is the Ohio Valley, where Rafinesque created numerous
new “species.” Another is the St. Lawrence Valley and northern New York.

This was followed by a presentation by Mr. F. R. Swift on “Treating Yeast Plants as
Individuals,” illustrated with splendid motion pictures.

In the latter, glass cover-slips are pre-coated with a vegetable-mineral oil mix-
ture, adjusted to fit the medium in use at the time. Small hanging drops are then
distributed on the cover-slips and each one is seeded with one yeast cell. It was ex-
plained that by varying the proportions of the vegetable and mineral oil with the
varying surface tension of different media being used: easily handled, uniform drop-
lets, can be assured.

The development of yeast cultures growing and sporulating in such droplets was
shown in a series of slides and by stopmotion photography, in a motion picture.

The discussion of these papers was continued after the meeting was formally adjourned
at 5:05 p.m., while tea was generously provided by The New York Botanical Garden.

Epwin B. Matzke, CoRRESPONDING SECRETARY.

FesruARY 2. MEETING IN THE MusEUM OF NATURAL HIsTorRY.

The meeting was called to order by the President, Dr. Robbins, at 8:15 p.m. At-
tendance 43. The minutes of the preceding meeting were accepted. Five new members
were elected to Annual membership. Dr. Seaver reported that the Auditing Committee
had found the Treasurer’s books in excellent condition. The report was accepted. Pres-
ident Robbins then read the names of those appointed to the various standing committees
of the Club. The scientific program was presented by Mr. G. L. Wittrock of The New
York Botanical Garden who spoke on “Local Plants Used by the American Indians,”
and illustrated these with colored slides. After a discussion period the meeting ad-
jyourned at 9:40 p.m.

Honor M. HottrncHurRst, RECORDING SECRETARY.

FEBRUARY 17. MEETING AT THE NEW York BOTANICAL GARDEN.

80 i ORR VAR.

program was Dr. Frances E. Wynne who spoke on “Variability and Distribution of
Drepanocladus in North America.”

Drepanocladus, like many aquatic and semi-aquatic plants, is extremely variable.
Field and herbarium studies have been made to determine which variations are
hereditary and which are merely environmental fluctuations. Careful examination oz
leaves irom different parts of the same plant shows that elongated leaves, costae,
and cells are always produced when the plant grows submerged in water, whereas
shorter leaves and cells are produced by stems which grow emergent. Many of the
described varieties are merely seasonal phases produced by changes in the water
level. The present monographic study has reduced the previously recognized 24 spe-
cies and 30 varieties to 9 species, 1 subspecies, and 4 varieties. Hereditary factors
determine the presence of an excurrent costa and secund leaves; therefore these
characters are used as the basis for varieties in several species. The environmental
fluctuations of the shape of the leaves, costae, and cells are not given taxonomic
recognition.

The species of Drepanocladus may be divided into two categories on the basis
of fundamental variability. All the boreal-montane are extremely adaptable ane
variable and as a result of their toleration of a large variety of habitats have spread
over a wide range. The arctic-alpine species are stable, clear-cut species limited to
one region and one type of habitat. |

Drepanocladus has a circumpolar distribution in both hemispheres. In North
America its present range coincides with the maximum extent of continental and
cordilleran glaciation during the Pleistocene. In eastern North America the distribu-
tion in partially glaciated states such-as Pennsylvania, New Jersey, Ohio, Indiana,
and Missouri is significant. In these states Drepanocladus does not occur south of
the till sheets except in a few isolated stations. In western North America it is found
in mountain bogs and alpine meadows.

Two types of distribution result from the Pleistocene glaciation: (1) relic, static
and (2) general, widespread. Any hypothesis, proposed to explain the post-Pleisto-
cene dispersal of plants, must consider these two types of distribution found on
glaciated areas. Of the numerous explanations which have been proposed, the most
satisfactory is founded on the genetic constitution of the plants. Species may be
plastic and adaptable or rigid and static. The boreal-montane species of Drepanoc-
ladus are adaptable because a large number of individuals survived the Pleistocene
in a large variety of habitats on all of the possible refuges; therefore a large number
of biotypes contributed to these plastic species. The arctic-alpine species are rigid
because only a few individuals survived in a few habitats on only one of the refuges;
the biotypes contributing to these species were depleted by the vicissitudes of the
ice age leaving the species genetically rigid.

The second speaker was Dr. Morris Winokur who spoke on “Photosynthesis in Bac-
Peticen

The attempt to interpret the metabolism of the green and sulfur bacteria has
resulted in the development of a generalized concept of photosynthesis which is
applicable to the green plant as well.

ING AMINIMITIIES, (OUR Wills: EIOGs: 81

use of different biological materials: Engelmann—purple sulfur bacteria; Molisch—
purple non-sulfur bacteria; Wuinogradsky—colorless sulfur bacteria. Buder at-
tempted to harmonize the diverse views by categorizing the organisms employed.
The existence of an intimate connection between the photosynthetic activity of the
purple sulfur bacteria and their respiratory phenomena was first clearly expressed
by Kluyver and Donker.

The consequence of the acceptance of this broad generalization is that it renders
untenable the classical Willstatter-Stoll theory of green plant photosynthesis.

After discussion of both papers, the meeting adiourned at 4:35 p.m., to be continued
over the inviting tea and refreshments served by friends at the Garden.

Honor M. HoLtitincHurst, RECORDING SECRETARY.
Marcu 2. MEETING IN SCHERMERHORN EXTENSION, COLUMBIA UNIVERSITY.

The fungus or fungi which induce mycorrhiza formation on the roots of shortleat
pine in the Missouri Ozarks area are apparently obligate parasites. The nature of
the symbiotic relationship between fungus and tree roots is determined largely by
environmental factors. When the trees are grown upon soils which are nutritionally
poor or unbalanced or when light intensity is low, fungal invasion of the short roots
readily occurs, and many well developed mycorrhizae are formed. Tree growth may ~
vary from slow to negligible. When the trees are grown upon fertile soil mycorrhiza
formation is difficult and slow and tree growth may be good, but if the soil contains
appreciable amounts of organic matter the seedlings will most likey fall victim of
damping-off fungi. When the trees are grown upon soils which contain relatively
low amounts of the necessary nutrients but when these nutrients are present in
physiologically balanced proportions on the soil colloids mycorrhizal development
and tree growth are both favored. Under such circumstances the mechanism of the
beneficial effect of mycorrhizae upon tree growth seems to be associated with an
increased salt absorption which is conditioned by an increased rate of aerobic respira-
tion and by a newly introduced mechanism for anaerobic respiration both of which
serve to maintain the energy output necessary for the growth processes. Hydrogen
ion excretion by the roots under such circumstances is increased several times thus
making it possible for the root colloids to undergo more intensively base exchange
reactions with the soil colloids in the initial phase of salt absorption. The enhanced
absorption of salts may then bring about greater water absorption and resulting in-
creases in both volume and mass of the plant tissues.

Following the discussion period, the meeting adjourned at 9:35 p.m.
Honor M. HotiincHurst, RECORDING SECRETARY.

82 TORRE AYA

Marcu 17. Meetine at THe New York BoranicaL GARDEN.

The meeting was called to order by the Vice-President, Dr. Seaver, at 3:30 p.m.
Attendance 36. The minutes of the preceding meeting were accepted. Dr. Whaley an-
nounced the death of Dr. Tracy Hazen on March 16th, in Waterbury, Conn. On a
motion by Dr. Stewart it was voted that the Secretary send the condolences of the Club
to the family of Dr. Hazen. Dr. Matzke stated that he had received from Dr. Moulton,
of the American Association for the Advancement of Science, a request for a summary
of the history of the Torrey Botanical Club. Dr. Matzke said he would be willing to
prepare this summary which is to be published in the Journal of the Association with
the histories of other affiliated societies. The first scientific paper was presented by Dr.
Ernest Naylor who spoke on “Problems of Cellular Behavior during Regeneration.”

The author presents a brief discussion of some of the cell changes during early
stages of shoot and root formation on isolated plant parts during regeneration. The
multiplication and organization of cells during regeneration involves two funda-
mental types of cells morphologically. One is the meristematic type, which may or
may not be definitely organized into recognizable growing points. Such cells may be
variously located in leaf axils, nodal regions, leaf margins, woody structures, and
in other places.

The other type is concerned with differentiated cells of the plant body which
undergo structural changes and become actively meristematic to produce the new
root and: shoot primordia. Such de-differentiation of vacuolate cells is described in
various tissues of a number of seed plants. The extent and limitations of such de-
differentiations in plant cells is briefly considered and some of the theoretical im-
plications pointed out.

Dr. Whaley was the second speaker on the scientific program, and his topic was “In-
feriority Complexes in Plants.”

Recent work of Dobzhansky and others indicates that in natural populations
many detrimental recessive genes are accumulated. The number and relative potency
of these genes is dependent upon the population structure, which is a function of the
number of individuals and the type of reproductive mechanism. Under selection it is
also possible for unfavorable dominants to accumulate. Heterosis is the result oi
masking of these deleterious recessives in some organisms, the result of heterozy-
gosity in others. Suggestions as to the nature of some of these deleterious factors
is found in excised root culture experiments. The roots of certain tomato lines show
a deficiency in ability to synthesize pyridoxine, others in the ability to synthesize
nicotinamide. Crosses between such lines produced vigorous hybrids under ordinary
field conditions. Hybrid vigor represents a return to an “optimum” phenotype rather
than any “super” phenotype.

After a discussion of both papers, the meeting adjourned at 4:35 p.m. Then tea and
delicious refreshments, in keeping with the spirit of St. Patrick’s Day, were served by
friends at the Garden.

Honor M. HoLtitincHurst, RECORDING SECRETARY.

Marcu 27. Fietp Trip to The New York Zoological Park for the study of some animal
habits. Leader, Miss Nellie L. Condon, Director, Reptile Study Society of America.
Attendance 11.

Marcu 28. Fietp Trip to Springdale, N. J., for limestone lichens. Leader, Mr. G. G.
Nearing. Attendance 4. Unusual forms found were: Acarospora murorum, Cypheluun
tigillare, and Physcia venusta. The last two were in fruit, and these fruiting forms
appear to be rare.

Aprit 4. Fretp Trip to Central Park, N. Y. to search for the trees mentioned in L. H.
Peet’s book “Trees and Shrubs of Central Park’ (1903). Leader, Dr. E. B. Matzke,
Columbia University. Attendance 25. Many of those present took an active part in the

ANC IMIWALIINES OU Wists, CILIU18} 83

identification of the trees, and all of us profited by Mr. James Murphy’s generous and
genial contributions on the trees and shrubs as well as on the history and lore of Central
Park. The following plants were found in flower: Cornus mas, Lonicera fragrantis-
suma, Ulmus americana, U. campestris, and Acer rubrum. Most of the day was spent
trying to locate trees mapped in Peet’s book. The morning was devoted to the southern
half of the park, and the afternoon to the northern end. A few tentative conclusions may
be suggested, subject of course te correction after more careful study:

3. The Turkey Oak, Quercus Cerris, has grown and perhaps prospered; native
oaks apparently do not thrive.

4. The English Elm, Ulimus campestris, seems to have done reasonably well,
distinctly better than our native ones.

5. In the wetter habitats the red maple, Acer rubrum, seems to be pretty well
established.

6. Ailanthus apparently “seeds in” in the park.

Epwin B. Matzke

ApriL 6. MEETING IN SCHERMERHORN HALL, CoLuMBIA UNIVERSITY.

Dr. Matzke then read to the club the letter which he, as Corresponding Secretary,
had sent to the family of the late Dr. Tracy E. Hazen:

New York City
March 23, 1943
Dr. Ropert HAZEN
Thomaston
Connecticut.
Dear Dr. Hazen:

At its meeting held on March 17, 1943, the Torrey Botanical Club directed its
Secretary to extend sympathy and condolence to the family of the late Professor
Tracy Elliot Hazen.

Your grief, and ours, may be assuaged by a knowledge of Professor Hazen’s
goodness, of his quiet nobility, and of his high attainments.

The Torrey Botanical Club realizes that a faithful officer, member, and friend
has passed to his reward; it is grateful for having shared in the innate richness of
his life.

In deep respect,
Epwin B. Matzke,
CORRESPONDING SECRETARY.

iva)
+e

TORREYA-:

Dr. Robbins announced that he had appointed a committee to drait a biographical note
on Dr. Hazen for the BULLETIN. The committee consists of Dr. Carey, chairman, Dr.
Barnhart, and Dr. Bold.

The scientific program oi the evening was presented by Dr. A. B. Stout of The
New York Botanical Garden, who spoke on “Dichogamy in Relation to Reproduction,”
illustrated with lantern slides. Aiter questions and discussion from the floor, the meet-
ing adjourned at 9:40 p.m.

Honor M. HoLt_tincHursT, RECORDING SECRETARY.

Aprit 11. Fretp Trip to The Brooklyn Botanic Garden and Conservatories for seasonal
studies outside, and for observation of economic plants from other lands. Leaders,
Dr. A. H. Graves and Dr. A. Gundersen of the Garden staff. Attendance: 3 members
and 147 visitors in the Garden.

Aprit 17. Fretp Trip to The New York Botanical Garden Conservatories, particularly
the Easter exhibit in the Display House featuring trees of the Holy Land. Leader, Dr-
H. A. Gleason of the Garden staff. Attendance 11.

Aprit 18. Fretp Trip to the Lichen Trail in Palisades Interstate Park for lichens, fungi,
and general botany of the season. Leader, Mr. G. G. Nearing. Attendance 4.

Aprit 21. MEETING AT THE BROOKLYN BoTANIC GARDEN.

The meeting was called to order at 3:30 p.m. by Dr. C. Stuart Gager, in the absence
of the President and Vice-Presidents of the Club. Attendance 18. The minutes of the
preceding meeting were approved. The program consisted of a talk by Dr. Henry K.
Svenson on the “Plants of a Long Island Pond,” illustrated with Kodachrome slides ;
and an inspection of the Local Flora Area of the Garden under the leadership of Dr-
Svenson.

Honor M. HoLiincHurRST, RECORDING SECRETARY.

Aprit 24. Frerp Trip to Surprise Lake, Watchung Reservation, near Summit, N. J., ior

reptiles, amphibia, and spring plant life, all of which reflected the late season. Leader, —

Miss Nellie L. Condon. Attendance 11].

May 1. Fretp Tri to Mertensia Island along Raritan River above Raritan, N. J., to see
the profuse stand of Mertensia, Dentaria, Erythronium, etc. This was the ideal date ior
this season. Leader, Dr. John A. Small, New Jersey College for Women. Attendance 6.

May 2. Fretp Trip to Silver Lake, White Plains, N. Y., for spring flowers and birds.
The day was cold and windy. 25 bird species were seen, 4 violets and 10 other plant
species were found in bloom. Leader, Miss Farida A. Wiley, American Museum of
Natural History. Attendance 12.

May 8-9. WEEK END FieLp Trip to Camp Thendara, Lake Tiorati, Palisades Interstate
Park, N. Y., for study of birds and plants. Leader, Mrs. Richard M. Abbott. Attendance
32, of which at least 5 were from the Torrey Club. 69 bird speceis were recorded,
including 19 warblers. :

May 11. MEETING 1n SCHERMERHORN Hatt, CoLUMBIA UNIVERSITY.

The results already obtained from-the systematic studies of woods indicate

clearly that any wood sample is identifiable. The unit of classification is at present
the genus, but well-defined species are frequently recognizable and their number

IMC II WINNS Ss) Ole Mave, (CLIO) 85

will increase with fuller knowledge of their range of variation. Ability to identify
a wood is of practical value to timber dealers and users and an important aid to
taxonomists in determining imperfect herbarium material and in preventing or cor-
recting faulty classification.
The essentials for systematic study of woods are: 1. A comprehensive and repre-
sentative collection of samples obtained with herbarium material ‘determined by
competent taxonomists. In the Yale collections there are 40,700 catalogued samples
representing nearly 12,000 named species of 2,800 genera and 232 families. 2. A col-
lection of slides with cross, radial, and tangential sections for examination under the
microscope. The Yale slide collection contains about 19,500 slides of 11,072 specimens |
of 6,506 name species, 2,616 genera, and 218 families. 3. Careful examination of the
slides by trained anatomists and the preparation of descriptions and tabulations of
all essential features. The standards used are those approved by the International
Association of Wood Anatomists after several years of cooperative effort. 4. The
use of the assembled data for making keys or other aids to identification. Numerous
keys to special groups have already been published and others are in preparation.
The task is very large, difficult, and costly and can only be carried out success-
fully through cooperative efforts. Ordinary taxonomists, though willing to accept
the aid of the anatomist in a time of trouble, make no effort to secure material
essential for anatomical study. Fortunately there are exceptions to this rule and
The New York Botanical Garden is foremost among American institutions in en-
couraging its botanists to collect wood samples. Systematic wood anatomy has made
its greatest progress during the past decade for the simple reason that during that
time research workers in various parts of the world effected an organization and
pooled their efforts and materials. The best incentive to further progress would be
the addition of new and better material which botanical expeditions could so readily
supply.
Because discussion of the talk was sharply curtailed at 9:30 p.m. by the sounding of
sirens for an air raid drill, the meeting quickly adjourned to darkened halls, where by
the light of a lantern, Dr. Record graciously identified wood specimens presented by
members of the audience.
Honor M. HoLtLtincHursT, RECORDING SECRETARY.

May 15. Fietp Trip to McLean Woods, The Bronx, N. Y., for spring study of the area.
Species lists were prepared and filed with the Field Committee. Leader, Mrs. Mary
Holtzoff. Attendance 14.

May 16. Fietp Trip to Point Pleasant, N. J., to search for Britton’s Violet, of which a
good stand was found, and in addition a large number of other plants. Leader, Mr. Louis
Hand. Attendance 7.

May 21-23. Fretp Trip to Culvers Lake, N. J., for the Annual Branchville Nature Con-
ference. In order to have the conference at the most desirable season and without in-
creased expense to those participating it was necessary to change from THE PINES
to THE HALTERE for accommodations. This proved satisfactory. Leaders: Mr.
Wallace M. Husk, Professor Oliver P. Medsger, and Dr. Julius Johnson. Attendance
40.

May 22. FieLp Trip to Ridgewood, N. J., to see the Rhododendron seedbeds, nurseries, and
stock of the leader, Mr. G. G. Nearing. Attendance 10.

May 29. Frétp Trip to Haskell, N. J., for fungi of the Chicohikie Falls region, especially
Fissipes acaulis, of which there was plenty. Leader, Mr. F. R. Lewis. Attendance 5

= =T ies r
7 - f= 1

THE TORREY BOTANICAL CLUB

Council for 1943
Ex officio Members

William J. Robbins Lela V. Barton Harold W. Rickett
C. Stuart Gager Edwin B. Matzke Michael Levine
John S. Karling Honor M. Hollinghurst John A. Small
Fred J. Seaver W. Gordon Whaley Bernard O. Dodge

Elected Members

1941-1943 1942-1944 1943-1945
John H. Barnhart : J. M. Arthur Charles A. Berger
R. C. Benedict W. J. Bonisteel Clyde Chandler
Helen M. Trelease Arthur H. Graves Albert E. Hitchcock
P. W. Zimmerman Sam F. Trelease Roger P. Wodehouse

Committees for 1943
ENDOWMENT COMMITTEE

Clarence Lewis, Chairman Henry de la Montagne
J. Ashton Allis Helen M. Trelease
Caroline C. Haynes

PROGRAM CoMMITTEE

Edwin B. Matke, Chairman (ex officio) A. B. Stout
Charles A. Berger W. Gordon Whaley
Arthur H. Graves P. W. Zimmerman ve

Honor M. Hollinghurst

Fietp CoMMITTEE

Joun A. SMA, Chairman

Edward J. Alexander Inez M. Haring Rutherford Platt
Vernon L. Frazee Michael Levine Daniel Smiley, Jr.
Eleanor Friend H. N. Moldenke Henry K. Svenson
Alfred Gundersen James Murphy Farida A. Wiley
Robert Hagelstein G. G. Nearing Ellys B. Wodehouse

LocaL FLtora COMMITTEE

Epwin B. Martzxe, Chairman

William J. Bonisteel Dolores Fay Harold W. Rickett

Herbert M. Denslow John M. Fogg, Jr. Hester M. Rusk

James Edwards H. Allan Gleason Ora B. Smith
Cryptogams

Ferns and Fern Allies: R. C. Benedict, W. Herbert Dole, N. E. Pfeiffer
Mosses: E. B. Bartram

Liverworts: A. W. Evans, E. B. Matzke

Freshwater Algae: H. C. Bold

Marine Algae: J. J. Copeland

Fungi: A. H. Graves, J. S. Karling

Lichens: J. W. Thomson, Jr.

Mysxomycetes: R. Hagelstein

PuBLICATIONS EXCHANGE COMMITTEE

Edwin B. Matzke, Chairman (e% officio) Amy L. Hepburn Lazella Schwarten

OTHER PUBLICATIONS
OF THE e 5

TORREY BOTANICAL CLUB

lished bi-monthly at present. Vol. 69, published in 1942, contained 711

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In addition to papers giving the results of research, each issue con-

tains the INDEX TO AMERICAN BoTANICAL LITERATURE—a very compre-

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Of former volumes, 24-69 can be supplied separately at $6.00 each;
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(75 cents) will be furnished only when not breaking complete volumes. ©

(2) MEMOIRS +
The Memoirs, established 1889, are published at irregular intervals.

Volumes 1-18 are now completed. Volume 17, containing Proceedings

of the Semi-Centennial Anniversary of the Club, 490 pages, was issued
in 1918, price $5.00. EY
Volume 18, no. 1, 108 pages, 1931, price $2.00. Volume 18, no. 2,
220 pages, 1932, price $4.00. Volume 18 complete, price $5.00. gz
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(3) INDEX TO AMERICAN BOTANICAL LITERATURE

Reprinted monthly on cards, and furnished to subscribers at a *
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Columbia University,
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PLT arian |

; NEW

Volume 43 December 1943 Number 2po7;
GA
EDITED FOR
THE TORREY BOTANICAL CLUB
BY

HAROLD H. CLUM

John Torrey, 1796-1873

CONTENTS

THE SEVENTY-FIFTH ANNIVERSARY CELEBRATION OF THE TORREY
BOTANICAL CLUB, JUNE 22-27, 1942

Viruses in Relation to the Growth of Plants...................... L. O. KunKeL 87
Animal Hormones Affecting Growth and the Several Effects of :

SInSLEGEIOLMONES 237 ody lai steed a ede cree a cuhefeds Se elslelateretatere en Oscar RIDDLE 96
The Formative Influences and Comparative Effectiveness of

Various Plant Hormone-like Compounds.................. P. W. ZIMMERMAN 98
Plants Need Vitamins Too....................0ee eee eeeee Wuu1am J. Rossins 116
Genetics, the Unifying Science in Biology...................-. GerorceE H. SHULL 126
Criteria for the Indication of Center of Origin in Plant

Geographical Studies. 20. 20.5.1. 0. 2505 Ss eae ea Stantey A. Carn 132
Phytopathology—1867-1942.. 1.2... 0.0... c cee cee ee tee GrorceE M. Reep 155

The Field Trip to the New Jersey Coast and Pine Barrens,
Friday and Saturday, June 26-27, 1942....E. J. ALEXANDER AND H. K. Svenson 170

Activities of the Club, May to November 1943...............20 0c. c eee eee eens 174
Additions to the List of Botanists in the Frontispiece.....................0200e002: 178
MNES! £0: CORREYA : V OLIMME ABs) s si2 uaa ois hel av eave tar lars win elmtatal aye] alas a ard ath ere racal dharat ef lem 179

PUBLISHED FOR THE CLUB

By tHe Free Press PrinTING ComMPANy
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Entered as second class matter at the post office at Burlington, Vermont,
October 14, 1939, under the Act of March 3, 1879

a 5 4 fs

THE TORREY BOTANICAL CLUB
OFFICERS FOR 1943

President: WILLIAM J. RopBins

Ist Vice-President: FRED J. SEAVER Recording Secretary: Honor M. Hot- ‘
2nd Vice-President: LELA V. BaRToN LINGHURST
Corresponding Secretary: EDWIN B. MATZKE Treasurer: W. GorpoN WHALEY

Editor: Harotp W. RicKETT

Associate Editors:

Irvinc W. BAILey ADRIANCE S. FOSTER
Epwarp W. BERRY Henry A. GLEASON
STANLEY A. CAIN ARTHUR H. GRAVES
M. A. CHRYSLER JouHn W. SHIVE
Harotp H. CLum R. P. WodEHOUSE

MICHAEL LEVINE
Business Manager: MicHAEL LEVINE Bibliographer: Mrs. LazELLA SCHWARTEN 4
Delegate to the Council, N. Y. Academy of Sciences: BERNARD O. DoncE

Representatives on the Council of the American Association for the
Advancement of Science

Joun H. BARNHARDT ALBERT F, BLAKESLEE

Representative on the Board of Managers of the N. Y. Botanical Garden:
Henry A. GLEASON

MEMBERSHIP IN THE TORREY BOTANICAL CLUB

All persons interested in botany are invited to join the club. There are four classes ©
-of membership: Sustaining, at $15.00 a year; Life, at $100.00; Annual, at $5.00 a year
and Associate, at $2.00 a year. The privileges of members, except Associate, are: (a) To ©
attend all meetings of the club and to take part in the business, and (b) to receive its —
publications. Associate members have the privilege of attending meetings, field trips
and of receiving the Schedule of the Field Trips and the Bulletin of the New York —

Academy of Sciences.
TORREYA

TorREYA was established in 1901 as a monthly publication of the Torrey Botanical Club
for shorter papers and interesting notes on the local flora range of the Club. It also ©
contains the proceedings of the Club, reports of field trips, and some book reviews and
news notes. The Council of the Torrey Botanical Club has decided to devote volume 43 —
of TorreyA, 1943, to the publication of the papers presented in June 1942 at the 75th Anni- —
versary Celebration of the Club, and to the Proceedings of the Club. This volume will
be published in two numbers. y

TorREYA is furnished to subscribers in the United States and Canada for one dollar
per year (January-December): single copies thirty cents. To subscribers elsewhere, —
twenty-five cents extra, or the equivalent thereof. Postal or express money orders, drafts, —
and personal checks are accepted in payment. Subscriptions are received only for full ©
volumes. ’

Claims for missing numbers should be made within sixty days following their date ~
of mailing. Missing numbers will be supplied free only when they have been lost in the ©
mails. All subscriptions and requests for back numbers should be addressed to the treasurer, —
Dr. W. Gordon Whaley, Barnard College, Columbia University, New York, N. Y. 4

Of the annual membership dues of the Torrey Botanical Club, $.50 is for a year’s
subscription to TORREYA.

TorREYA is edited for the Torrey Botanical Club by 3

HAROLD H. CLUM
HUNTER COLLEGE, 695 ParK AVENUE
New York, N. Y.

TORREYA

Vout. 43 DECEMBER 1943 No.

Viruses in Relation to the Growth of Plants*

L. O. KUNKEL

About twenty-four years ago, Nishimura (10) reported that Physalis
alkekengi allowed the tobacco mosaic virus to multiply within its tissues but
showed no symptoms of disease. Since that time other masked carriers of
plant viruses have been studied (2, 3, 9). We now know that practically all
potatoes produced in this country carry the X virus but that, unless it occurs
in combination with some other potato virus, no well defined symptoms are
produced (3). We also know that some of the mutants of ordinary tobacco
mosaic virus cause no obvious symptoms in tobacco (2). But, while it is true
that some viruses multiply in some plants without causing symptoms by which
a disease can be readily recognized, it is doubtful whether there are any really
symptomless carriers. All viruses that become systemic and multiply within
a plant probably cause some injury. However, the injury may be slight and
easily overlooked unless control plants are available for comparison. Some
virologists have gone so far as to suggest that there may be viruses capable
of stimulating rate of growth in plants, but if such viruses exist they have not
been discovered.

From viruses that cause exceedingly mild diseases, it is possible to pass
by gradual steps to viruses that are lethal. We may, in fact, do this without
going outside of the tobacco mosaic virus group. When masked strains of
tobacco mosaic virus are propagated in tobacco, they are sooner or later re-
placed by mild mottling strains some of which approach ordinary tobacco
mosaic virus in severity. Similarly, when severe strains are propagated in
tobacco, they are replaced by milder strains some of which approach tobacco
mosaic virus in mildness. All except the so-called masked virus strains cause
marked stunting and other symptoms of disease. The masked virus strains
cause stunting but no other well marked symptoms. Thus, the tobacco mosaic
viruses and all other plant viruses may be classified as growth-depressing en-
tities. This, however, does not mean that they depress rate of growth in all
tissues.

The Fiji disease virus of sugarcane causes well marked galls in phloem
tissues (4). The cranberry false blossom virus, with which the writer has

* Read at the 75th Anniversary Celebration of the Torrey Botanical Club at the Boyce
Thompson Institute for Plant Research, Inc., Wednesday, June 24, 1942.

TorreyA for December (Vol. 43, 87-183) was issued February 10, 1944.
87

88 TORREYA

been working recently, causes increased growth in flowers. It depresses growth
in the plant as a whole but stimulates rate of growth in flowers. In a number
of different plants to which it has been taken, it causes the production of giant
blossoms. Its action on tomato flowers is shown in figure 1. In the truss on
the left the flowers are normal, while in the other two they are diseased. The
sepals of the diseased flowers are much larger than sepals of normal flowers.
Instead of remaining separate as in healthy blossoms, the affected sepals have
fused to form a sac-shaped structure. When the sac was torn open, it was
found that the petals were green in color and borne at the end of a thick stalk
which was about an inch in length. The petals were leaf-like in structure; some

Fic. 1. False blossom in tomato. The flowering truss at the left is healthy; the other
two are diseased. (Photograph by J. A. Carlile.)

were simple and others compound. The anthers were usually small and green.
In its effects on tomato flowers, false blossom resembles the big-bud disease
which occurs in the western part of this country (1) and in Australia (11).
Other effects of false blossom on flower trusses are shown in figures 2 and 3.
The diseased truss pictured in figure 2 was about four times as long as the
healthy truss. It terminated in two stem tips bearing leaves. The stem of the
diseased truss also was thicker than that of the healthy truss. Figure 3 shows
other variations in the deformation of enlarged flower trusses.

Some diseased trusses that had not elongated so much but were borne on
thick stems are displayed in figure 4 beside a normal truss. This type of mal-
formation was met with less often than the big-bud type. Apparentiy there are
several different strains of false blossom virus prevalent in nature. From some
diseased cranberry plants a strain was obtained that caused a severe check to
longitudinal growth but stimulated transverse growth. Plants with this strain
stopped producing flower buds soon after they were infected and did not stimu-
late the production of secondary shoots. From other false blossom cranberry

KUNKEL: VIRUSES

Fic. 2. False blossom in tomato. The flowering truss at the right is healthy; that at the
left diseased. The picture shows the stimulating effect of the virus on flowering branches.

Fic. 3. False blossom in tomato. The three flowering trusses show different effects of the
virus. (Photographs by J. A. Carlile.)

TiO RARE Yue

Fic. 4. False blossom in tomato. The flowering truss at the left is healthy ; the other
three are diseased, showing thickening of stems.

Fic. 5. False blossom in tomato. In the tip at the left longitudinal growth has been
stimulated and transverse growth checked, while in the tip at the right longitudinal growth
has been checked and transverse growth stimulated. The other three tips show interme-
diate effects. (Photographs by J. A. Carlile.)

KUNKEL: VIRUSES 91

plants a strain was obtained that stimulated longitudinal growth and checked
transverse growth. This caused a spindling witches’ broom type of growth. In
tomatoes with the spindling strain no flower buds were produced except shortly
after infection. The flowers that were produced usually were not more than two
to three times the size of normal flowers. Between these extremes in which long-
itudinal growth was almost entirely stopped but transverse growth stimulated,
on the one hand, and in which longitudinal growth was greatly stimulated but
transverse growth severely checked, on the other hand, were strains that
caused intermediate effects. Some of these are shown in figure 5 where tips
from five different diseased tomato plants are pictured. The tip at the extreme
right 1s greatly shortened and thickened; that on the extreme left is tall and
spindly. The three types shown between these exhibited intermediate ef-
fects. The virus obtained from most diseased cranberry plants caused the
symptoms shown by the tip in the center of the picture. This is the typical
big-bud type of top where flowers are large and malformed and where con-
siderable numbers of secondary shoots are produced. When scions from plants
affected in this way and scions from plants showing the two extreme effects
were grafted to healthy tomato plants, each came down with the type of disease
characteristic of that shown by the plant from which the scion was taken. When
the two extreme types were transmitted to periwinkle plants, they caused simi-
lar variations in symptoms. The virus that depressed longitudinal growth but
stimulated transverse growth in the tomato caused the production of short thick
tips but very little chlorosis or stunting of leaves when taken to periwinkles.
The virus that stimulated longitudinal growth but depressed transverse growth
in stems of the tomato produced similar effects in stems of periwinkles. In
leaves it caused a marked chlorosis and narrowing. When the virus causing
typical big-bud in tomato was taken to periwinkle, it caused the production of
green malformed flowers such as are shown beside normal flowers in figure 6.
When scions from periwinkles showing the different types of effects were
grafted to healthy periwinkles, each transmitted the disease characteristic of
the plant from which it was taken. While it has not been proved that these
different types of disorders are caused by strains of the cranberry false blossom
virus, this seems likely. When the common type of false blossom virus was
transmitted to the composite, Calendula, it caused the production of malformed
green flowers such as are pictured in figure 7 beside a healthy flower. Another
plant in which false blossom virus caused gigantism in flowers was Nicotiana
glutinosa. An early stage in the development of giant sepals is shown in figure
8, a late stage in figure 9. The blossoms at the left in both figures are normal ;
the others are diseased. Malformed leafy structures are shown protruding
from some of the diseased flowers in figure 9. At the same time that the virus
caused enlargement of flowers, it produced dwarfing of leaves.

TORRENA

Fic. 6. False blossom in Vinca rosea. The flowering branch at the left is healthy; that
at the right is diseased. The diseased flowers are malformed and virescent.

Fic. 7. False blossom in Calendula. The flower at the left is healthy ; the other two are
diseased. Affected flowers are green in color. (Photographs by J. A. Carlile.)

KUNKEL: VIRUSES

Fig. 8. False blossom in Nicotiana glutinosa. The flower at the left is healthy. The other
two flowers have false blossom.

Fic. 9. False blossom in Nicotiana glutinosa. The flowering branch at the left is healthy ;
that at the right diseased, showing gigantism in flowers affected by the virus. (Photo-
graphs by J. A. Carlile.)

94 DIO RERCE Nex

It would be possible to proceed at great length with the story of how false
blossom virus upsets growth relationships, enlarges, distorts, and malforms
flowers, and transforms plants of different species to such an extent that they
can scarcely be recognized. But there would be no point‘in doing this, for the
variability in the symptoms produced in different species is almost endless. It
perhaps is sufficient to say that in all plants to which false blossom virus was
taken it caused virescence and gigantism in flowers or parts of flowers,
chlorosis and dwarfing in leaves, and an elongating or a shortening of inter-
nodes of stems.

Another effect of certain viruses on growth in plants that needs to be men-
tioned is repression of dormancy and maturity. Many biennial and perennial
plants pass the winter in a dormant state. When brought into a greenhouse
where good growing conditions are maintained and where they might be ex-
pected to grow continuously, they become dormant or semi-dormant as the
winter season approaches. Many annual plants grow to maturity in a few
months and then die. When affected by certain virus diseases, perennial plants
fail to go into a dormant state regardless of environmental conditions. Peach
trees affected by yellows disease do not stop growing as cold weather comes
on but continue vegetative growth until the tender tips of branches are frozen
and killed.

China aster plants set in the field late in May or early in June blossom in
August and mature seeds in early autumn. By the time cold weather arrives,
all healthy plants have died. The course of events is very different for plants
that contract the aster yellows disease. They produce malformed virescent
flowers and sterile seeds (5). It is true that some diseased plants produce
viable seeds, but such seeds are borne only by flowers that have not been in-
vaded by the virus. Instead of affected plants maturing and dying as cool
weather approaches, they live and grow. They of course do not grow very fast
and are eventually killed by low temperatures. But there is a period during
which the only living plants in the field are those affected by the aster yellows
disease. It is thus clear that the virus lengthens the life of the plant.

When healthy potato plants are grown in greenhouses, they produce tubers,
mature, and die. If infected by the witches’ broom virus, they produce tubers
but they do not mature and die. Growth continues summer and winter for an
indefinite period of time. There is a potted potato plant with witches’ broom
in one of our greenhouses that has been growing there for more than two years.
Its healthy sister plants matured and died long ago. The witches’ broom virus
has had a favorable effect on the longevity of the plant.

Certain plant virus diseases are readily cured by heat. When affected plants
are held at moderately high temperatures for appropriate periods of time, the
viruses that cause these diseases are inactivated but the plants are not seriously

KUNKEL: VIRUSES 95

injured. Peach yellows (6), aster yellows (7), and witches’ broom of potato
can be cured by heat in certain of the plants they affect. The false blossom
disease also can be cured (8). When plants are cured, all of the bizarre effects
produced by the viruses causing these diseases disappear. Cured peach trees
become dormant when the season for dormancy arrives. Cured potato plants
live no longer than healthy plants. The stimulating and stunting effects of cran-
berry false blossom virus in periwinkles subside and disappear when the plants
are cured. It is apparent that these viruses affect growth only during the period
in which they are present in the plants. The malformations that develop while
plants are sick are of course not corrected by cure, but all new growth pro-
duced after plants are cured is normal. In this respect these viruses are like
the growth-promoting substances. Their effects do not outlast the periods
during which they are allowed to act but, unlike growth-promoting substances,
viruses cause a retardation of growth in all plants in which they are permitted
to multiply, although, as we have seen, they stimulate growth in certain tissues.
They also are entirely unlike growth-promoting substances in that they infect,
multiply, and cause disease.

, DEPARTMENT OF ANIMAL AND PLANT PATHOLOGY

THE ROCKEFELLER INSTITUTE FOF MEDICAL RESEARCH
PrIncETON, NEw JERSEY

Literature Cited

1. Dana, B. F. Occurrence of big bud of tomato in the Pacific Northwest. Phytopath.
30: 866-869. 1940.

2. Hotes, F. O. A masked strain of tobacco-mosaic virus. Phytopath. 24: 845-873. 1934.

3. Jones, L. K., ANpDERSON, E. J. & Burnett, G. The latent virus of potatoes. Phytopath.
Zeitschr. 7: 93-115. 1934.

4. KunkKEL, L. O. Histological and cytological studies of the Fiji disease of sugar cane.
Hawaiian Sugar Planters’ Assoc., Exp. Bull., Bot. Ser., 3: 99-107. 1924.

5. KuNKEL, L. O. Sterility caused by the aster yellows disease. Mem. Hort. Soc. N. Y. 3:
243-244. 1927.

6. KUNKEL, L. O. Heat treatments for the cure of yellows and other virus diseases of
peach. Phytopath. 26: 809-830. 1936.

7. KuNKEL, L. O. Heat cure of aster yellows in periwinkles. Am. Jour. Bot. 28: 761-769.
1941.

8. KUNKEL, L. O. False blossom in periwinkles and its cure by heat. Science 95: 252. 1942.

9. McWuorter, F. P. A latent virus of lily. Science 86: 179. 1937.

10. NisHimura, M. A carrier of the mosaic disease. Bull. Torrey Club 45: 219-233. 1918.

11. SAMUEL, G., BALp, J. G. & Earpiey, C. M. “Big bud,” a virus disease of the tomato.
Phytopath. 23: 641-653. 1933.

VoL. 43 TORREYA DECEMBER 1943

Animal Hormones Affecting Growth and the Several Effects of
Single Hormones*

Oscar RIDDLE

In higher vertebrate animals and man, the forms in which hormonal regu-
lation is best known, several hormones act as stimulants to growth. In some
cases this stimulus is fairly restricted or localized and only a single function or
special tissue is affected. But advancing information indicates that many
hormones affect a variety of functions and organs. For higher animals it has
been learned during the past 15 years that the center of hormonal regulation
resides in the anterior pituitary gland; its hormones may be called “trigger”
hormones. In large measure these “trigger’’ hormones stimulate other hormone
producing glands (thyroid, gonad, adrenal) whose products may thus in turn
be called “target” hormones (thyroxine, estrone, testosterone, cortin). Growth
processes are affected by both “trigger’’ and “target” hormones; one of the
former, prolactin, and one of the latter, estrone (or estrogens), are here util-
ized as illustrations of hormones which are not only related to growth but
which also exhibit a variety of actions.

Prolactin stimulates milk secretion in mammals and growth of crop-sacs
and production of crop-milk in pigeons. It sometimes reduces or prevents the
secretion of the “trigger” hormone, gonadotrophin. It releases broodiness
(fowl, pigeon) and maternal behavior (rats). Perhaps it prolongs the life of
corpus luteum cells, and stimulates their production of the hormone, proges-
terone. In pigeons, but not in rats, it seems to be the chief and best of hormones
for the promotion of bodily growth. It assists growth in dwarf mice and there
synergizes the action of thyrotrophin on growth. In pituitaryless pigeons pro-
lactin can increase body weight and intestinal and liver tissue to an extra-
ordinary degree, and likewise it can partially support the pancreas; but all
these actions can be shared or augmented by hormone of the adrenal cortex
(unpublished, Miller and Riddle), while still a third hormone, thyroxine, fur-
ther assists in maintaining the weight of the intestine and pancreas. It is a
moot question whether the pituitary gland produces a single “growth hor-
mone” or whether bodily growth (probably somewhat differently stimulated
in different species) is a summation of effects of various “trigger” and “tar-
get’’ hormones.

Estrone, or stable estrone-like substance, has been obtained from yeast,
rape seed, potatoes and female willow catkins—even from petroleum and lig-

* Presented in more detail at the 75th Anniversary Celebration of the Torrey Botanical
Club at the Boyce Thompson Institute for Plant Research, Inc. Wednesday, June 24, 1942.
Only an abstract of the discussion is published here.

tm ©

eT ee ene ee Se ee

ba, thins 4} ip epee te tte OO

RIDDLE: ANIMAL HORMONES 97

nite. Estrone produces localized growth in oviduct, mammary glands and
uterus. It reduces or suppresses the output of the pituitary hormone, gonado-
trophin. It affects bone development and general bodily form in some species.
It has an action on the calcium, phosphorus and fat of the blood. In the mental
sphere it affects sex behavior.

The past 15 years of study of the actions, interactions and automatic con-
trol of release of the pituitary hormones in the bodies of higher animals have
provided a purely natural basis for some of the most mysterious performances
and adjustments of our own bodies. Now, for the first time in the long history .
of man, human beings partly know a series of organs and substances which—
acting in high degree as a self-regulating system—largely control the fuller
expression of growth, the rhythms of reproduction, and some aspects of tem-
perament and behavior. In short, we have come to recognize our anterior
pituitary gland as the master or governing gland; also, the brain and this
master gland are now marked as the two truly basic sources of the strength
and competence of man. It should arouse biologist and layman alike to reflect
that up to our own time mankind has made its whole history—its conquests,
its arts, its literature, its laws, its religions, its philosophies—while wholly
ignorant of one of the two physical sources from which the abilities of an
individual human being are derived.

CARNEGIE INSTITUTION OF WASHINGTON

DEPARTMENT OF GENETICS
CoLtp Sprinc HaArsor, NEw YorK

Vot. 43 TORREYA DECEMBER 1943

The Formative Influences and Comparative Effectiveness of Various
Plant Hormone-like Compounds*

P. W. ZIMMERMAN

Plant physiology has gone a long way since Boysen-Jensen discovered that
the stimulus which causes cell elongation and tropic curvatures in coleoptiles
passed through a discontinuity of tissue and appeared to be of a chemical nature

- (1). Since that time many chemical compounds, natural and synthetic, have
been found which when applied to plants act like hormones. In addition to cell
elongation these substances cause cell division, induce new organs, prevent
abscission, inhibit buds, modify the pattern of organs, and otherwise regulate
the growth of plants. Such substances have been given various names as hor-
mones, auxins, growth substances, growth promoters, growth regulators, etc.
None of these designations is satisfactory because a single substance has the
capacity to induce several varied responses. The word “formative” has often
been used to describe the effects of hormone-like compounds on plants. This
term did not seem significant until recently when it was found that some of
these physiologically-active compounds have a decidedly regulating and “form-
ative’ effect on the new growths of the entire plant (5, 7, 3, 4). This is in con-
trast with locally induced cell elongation. The subject of this paper concerns
especially formative influences and comparative activity of several hormone-
like compounds which modify the pattern of leaves, flowers, and fruit and
which change the correlation phenomena of organs.

METHODS AND MATERIAL

The activity of growth substances was usually detected by curvatures re-
sulting from induced cell elongation or by formative effects on later growth.
The former response occurred within a comparatively short period of time
(20 to 60 minutes). Formative effects appeared in days or weeks after the
plant had time to produce new organs. The first evidence of formative effects
appeared on new leaves which were modified in size, shape, pattern, and tex-
ture. Later the effects appeared on flowers, fruit, growth habit, and SESE ES
phenomena of organs.

The chemicals were applied to plants in water solution, as lanolin prepara-
tions, and as vapors. Various spreaders were used with water solutions but were
generally considered not essential. Water solutions (10 to 300 mg./l.) were
sprayed on the plants with a nasal atomizer, applied to the soil, and injected
into the stem with a glass capillary tube. Lanolin (or other oily substance)

* Read at the 75th Anniversary Celebration of the Torrey Botanical Club at the Boyce
Thompson Institute for Plant Research, Inc. Wednesday, June 24, 1942.

ZIMMERMAN: FORMATIVE INFLUENCES 99

preparations were made up with a series of concentrations of the chemical,
ranging from 0.005 to 20.0 mg./g. of lanolin. These preparations were applied
to local parts of the plant with a giass rod. To induce epinasty the material was
applied to the upper side of a young leaf petiole ; to induce curvature of the stem
the material was applied along one side of a young stem. Chemicals active for
cell elongation caused negative (away from treated side) curvatures within a
short time. The same treatment also served to determine whether the chemicals
had a formative influence on new growth. Plants were exposed to vapors of
the various compounds under bell jars, closed greenhouses, glass cages, or
other closed containers which could be kept reasonably tight. The esters were
more volatile than acids or amides and were considered best for vapor treat-
ments. The ultimate effects, however, were the same for all. The amount of
ester required under the bell jar was less than 1 milligram. When heat was re-
quired to volatilize the chemical, a small amount was placed on a watch glass
which in turn was placed on a warm or hot inverted crucible under a bell jar.
In the greenhouse a hot plate supplied the heat and an electric fan circulated the
air. .

Most of the chemical compounds used in the experiments and listed in the
tables were synthesized in the Boyce Thompson Institute laboratories. A few
were available from commercial supply houses.

RESULTS

For the study of formative influences three groups of compounds stand
out above all others. They are 8-naphthoxy acids, substituted phenoxy deriv-
atives of the lower fatty acids, and substituted benzoic acids. The substituted
groups were nitro, amino, methyl, or halogen radicals. These were used alone or
in various combinations substituted in the naphthalene or benzene ring.

B-Naphthoxy compounds. B-Naphthoxyacetic acid and the higher homo-
logs, propionic and butyric acids, were the first observed to have special forma-
tive influences (5). They were found to have in common with other plant hor-
mone-like compounds the capacity to cause cell elongation, parthenocarpic de-
velopment of ovaries, and to induce roots. Table 1 shows a list of naphthoxy
and naphthalene compounds and their activity for cell elongation and formative
influences.

It is interesting to note that for cell elongation naphthoxy compounds must
have the chain of the molecule linked to the beta position in the ring while the
alpha position is required for naphthaleneacetic acid. a-Naphthoxyacetic acid
is inactive for cell elongation but has a slight formative influence. Neither a-
nor B-naphthaleneacetic acid has a formative influence which modifies the pat-

tern of leaves.

100 TORR EWA

TABLE 1. MoLtecuLar CONFIGURATION AND COMPARATIVE ACTIVITY OF NAPHTHOXY AND
NAPHTHALENE COMPOUNDS

Cell Modification of
Substances elongation tomato leaves
—O—CH2COOH
Active Active
$—Naphthoxyacetic acid
Ce
ome Active Slightly Active
a—(-Naphthoxy) propionic acid
a
—O—CHCOOH Active Active

a—($-Naphthoxy) -n-butyric acid

—O—CH2COOH

ee Inactive Slightly Active

a—Naphthoxyacetic acid

—CH2COOH
Gal Active Inactive
a—Naphthaleneacetic acid
pc ge a ht a a ll eae ae it Pan 8
—CH2COOH
Inactive Inactive
B—Naphthaleneacetic acid
ae COOH
Inactive Inactive

a—(a-Naphthalene) propionic acid

ZIMMERMAN: FORMATIVE INFLUENCES 101

Attention has been called to the fact that though 8-naphthoxyacetic acid
and its higher homologs have a formative influence on growth, there are quali-
tative differences in responses induced with these compounds (6). It should be
pointed out also that different species bring out further qualitative differences.
8-Naphthoxyacetic acid, for example, modifies the leaves of Turkish tobacco
(Nicotiana tabacum) while B-naphthoxypropionic acid has little or no effect
on the pattern of leaves of this species.

Substituted phenoxy compounds. There are many active substituted
phenoxy compounds. The activity appears to be related to the kind, number,
_and position of substituted groups in the benzene ring. In general, halogen sub-
stitutions bring about greater activity than methyl, amino, or nitro groups.
With a single halogen group substitution the para position is more effective
than the ortho. However, 2,4-dichlorophenoxyacetic acid is more effective than
either o0- or p-chloro phenoxyacetic acid (4, 7).

Nitro group substitutions do not activate the phenoxy molecule except in
the meta position. The amino group activates the molecule when substituted in
the para position. The chlorine atom in the ortho, meta, or para position acti-
vates the phenoxy molecule. These comparisons are shown in table 2.

Table 3 shows the comparative activity of non-substituted and chloro-sub-
stituted phenoxy compounds. Phenoxyacetic acid does not modify the pattern
of leaves though its propionic and butyric acid homologs do. This is in contrast
with ortho, para, and 2,4-dichlorophenoxy compounds where the acetic acid
form modifies leaves but the corresponding propionic and butyric acid homologs
do not. These are in further contrast with m-chlorophenoxy and 2,4,5-trichloro-
phenoxy compounds where neither the acetic acid forms nor their higher homo-
logs modify leaves though all are active for cell elongation. In all active phenoxy
compounds the nucleus of the molecule was linked to the chain at the alpha
carbon atom. Comparable beta linkages made inactive compounds.

2,4,6-Trichlorophenoxyacetic acid and the propionic acid homolog did not

cause cell elongation but had a slight capacity to modify organs. 2,3,4,6-Tetra-
chlorophenoxyacetic acid and 2,3,4,5,6-pentachlorophenoxyacetic acid were in-
active.
When the ortho and para positions were substituted with chlorine or a
methyl group making 2,4-dichlorophenoxyacetic or 2,4-dimethylphenoxyacetic
acids, the compounds were active for cell elongation and for modification of
leaves. The higher homologs, however, were different. a-(2,4-Dichlorophe-
noxy) propionic and butyric acid were not active for modification of organs
while the corresponding methyl-substituted compounds were active. This pe-
culiarity is illustrated in table 4.

The formative influence of 2,4-dichlorophenoxyacetic acid is illustrated in
figure 1 B and C. When the Nicandra physalodes plant on the right was ap-

102 y TORREYA

TABLE 2. THe DEPENDENCE FOR ACTIVITY UPON THE POSITION OF SUBSTITUTED GROUPS

CEE COOr GEScL oe ee
O O O
|
NOg2
NOs Y
Inactive Active NOs Inactive
CH2COOH Waeo ee can eer e
O 0 O
| ]
NH
NHo /
Inactive Inactive NH2 Active
CH2COOH CH2COOH rege re
O O :
i C1 I
Ci
Active Active Gi Active

proximately five inches high, it was sprayed with a water solution containing
12.5 mg. of the chemical per liter of water. All new growth thereafter produced
modified organs. The change in the pattern of leaves can be seen by comparing
the three old leaves near the base of the stem which were nearly mature when
the plant was treated with those of the new growth. Also the flowers of the new
growth were modified as shown in figure 1 B and C. Stems and leaves some-
times became fasciated and flowers appeared to arise from leaves (Fig. 1 C).
Calyx tubes often remain closed and prevented the corolla from emerging.
The veins of the leaves often crowded toward the midrib making a narrow —
leaf with only a ruffle of blade around the edge. The veins became nearly trans-
parent and the plants appeared to have a virus disease.

Substituted benzoic acids. Considered from the standpoint of a plant growth
substance, benzoic acid is an inactive compound. When, however, the nucleus
is substituted, the molecule may be activated. The degree of activity depends
upon the kind, number, and the position of substituted groups. Amino, nitro,

ZIMMERMAN: FORMATIVE INFLUENCES 103

TABLE 3. MoLEcuLAR CONFIGURATION AND COMPARATIVE ACTIVITY OF PHENOXY AND
SUBSTITUTED PHENOXxY COMPOUNDS

Cell elongation Modification
(epinasty ) of tomato leaves
threshold conc. threshold conc.
Substances mg./g. mg. /g.
ee
O
|
Phenoxyacetic acid 20 Inactive
Ce aa
O
|
a—(Phenoxy ) propionic acid 5 5
ene
O
|
a a—(Phenoxy)-n-butyric acid 5 5
eae i
‘
Cl
o-Chlorophenoxyacetic acid 1 0.25
ee Sanaa
i
Cl
a—(2-Chlorophenoxy ) propionic acid 1 Inactive
Se
i
Cl

a—(2-Chlorophenoxy) -n-butyric acid 1 Inactive

104 TORRE Vex

TABLE 3. (Continued)

Cell elongation Modification of

(epinasty ) tomato. leaves
threshold conc. threshold conc.
Substances mg./g. mg./g.
{ cae
O
|
m-Chlorophenoxyacetic acid 0.5 Inactive
Cil
Pe OOs
O
|
a—(3-Chlorophenoxy ) propionic acid 0.5 Inactive
ee he aie
O
|
a—(3-Chlorophenoxy)-n-butyric acid 0.5 Inactive
Cl :
eee
O
|
p-Chlorophenoxyacetic acid 0.25 0.06
Gi
Se eae
O
|
a—(4-Chlorophenoxy) propionic acid 0.5 Inactive
il
CE ik aaa
O
|
a—(4-Chlorophenoxy)-n-butyric acid i Inactive

ZIMMERMAN: FORMATIVE INFLUENCES 105

TABLE 3. (Concluded)

Cell elongation Modification of

(epinasty ) tomato leaves
threshold conc. threshold conc.
Substances mg. /g. mg. /g.
fence
I
Cl
2,4-Dichlorophenoxyacetic acid 0.015 0.003
Cl
CH Ae COOH
i
C1
a—(2,4-Dichlorophenoxy) propionic 0.5 Inactive
acid
Cl
raat
i
Cl
a—(2,4-Dichlorophenoxy ) -n-butyric 0.5 Inactive
acid
Cl
to 2»COOH
O
Cl
Cl 2,4,5-Trichlorophenoxyacetic acid 0.06 Inactive
Cl
a COOH.
a—(2,4,5-Trichlorophenoxy ) 0.03 Inactive
1 propionic acid
ee
o a—(2,4,5-Trichlorophenoxy ) -n- 0.1 Inactive
butyric acid
UN nei StI tlie

ZIMMERMAN: FORMATIVE INFLUENCES 107

and halogen groups appear to be the most important. Some substituted benzoic
acids have a pronounced formative influence on plants but have little or no
effect on cell elongation. One compound of the group, however, induced both
cell elongation and modification of leaves (3, 7). Table 5 shows a list of active
and inactive compounds.

Positions 2, 3, and 5 in the nucleus appeared to be the most important for
substitutions. For example, 2,3,5-triiodobenzoic acid and 2-chloro-3,5-diiodo-
benzoic acid had the most pronounced formative influence of any of the com-
pounds listed. In addition to modifying the pattern of leaves, they influence
flowering habit and correlation of organs (8). One to 5 mg. of triiodobenzoic
acid added to the soil of a 4-inch pot in which a tomato plant was growing was
sufficient to cause modification of growth of the stem, leaves, and flowers, to
cause axillary buds to grow flower clusters instead of the normal leafy shoots,
and to induce the terminal bud to terminate with a flower cluster instead of
continuing with a leafy shoot (Fig. 2 A and B). Similar results were obtained
by other methods of applying the chemical. It is effective as a lanolin prepara-
tion (1 to 10 mg./g. of lanolin), as a vapor applied in a closed container, and
as a water solution applied as a spray (25 to 100 mg./1.).

The results obtained with 2-chloro-3,5-diiodobenzoic acid were similar to
but even more striking than those described for triiodobenzoic acid. Both com-
pounds caused the terminal bud and axillary buds of tomatoes to grow flower
clusters instead of leafy shoots, but the individual flowers were different. Those
which grew under the influence of 2-chloro-3,5-diiodobenzoic acid were small
with inconspicuous petals and sepals supported with an abnormally stout pedun-
cle (Fig. 3 A). As the chemical influence became weaker and the plants began
to recover, large single flowers instead of clusters were produced irregularly
along the stem. The small flowers did not set fruit but the large ones func-
tioned as normal flowers (Fig. 3 A and B).

Another active substituted benzoic acid, 2-bromo-3-nitrobenzoic acid, is of
special interest since it caused both cell elongation (epinasty) and modified
leaves of tomato plants (7). It was not as active for cell elongation as some
of the phenoxy compounds but had a pronounced formative influence on growth.
2-Chloro-5-nitrobenzoic acid has a formative influence but does not cause cell
elongation. Judging from active benzoic acids listed in table 5 it would appear

Explanation of figure 1

Modification of organs induced with substituted phenoxy compounds. A. Tomato shoots:
left, control; right, response to spray with solution of B-(2,4,6-trichlorophenoxy ) -8’-chloro-
diethyl ether. B. Nicandra plants: left, control; right, modifications induced with 2,4-dich-
lorophenoxyacetic acid (12.5 mg./l.). Solution applied at tip with nasal atomizer when
plant was 5 inches in height. Note non-modified leaves at base which were present when
treated. C. Enlarged leaves, buds, and flowers taken from plants in B.

108 TO RAVE NEA

TABLE 4. VartaTIon in Activity ACCORDING TO THE POSITION OF CHLORO AND METHYL
SUBSTITUTED GROUPS

Cell Formative
Substances elongation effects
eee
O
(Cll
2,4-Dichlorophenoxyacetic acid Active Active
Cl
oo. haa OH
O
Cl
a—2,4-Dichlorophenoxy ) propionic Active Inactive
acid
Cl
See eae
t
Cl
a-(2,4-Dichlorophenoxy ) -1-butyric Active Inactive
acid
Cl
ingest
O e
CH3
2,4-Dimethylphenoxyacetic acid Active Active
CHs
CH ia COOH
O
CH3
a—(2,4-Dimethylphenoxy) propionic Active Active
acid
CHs3
oe
i
CHs3
a—(2,4-Dimethylphenoxy)-n-butyric Active Active
acid

CH3

i i i

ZIMMERMAN: FORMATIVE INFLUENCES

Fig. 2. Formative influence of 2,3,5-triiodobenzoic acid on tomato plants. A. Left,
control ; right, treated with 2 mg. of the chemical in 50 cc. of water applied to the soil when
the plant was approximately 5 inches in height. B. Left, control; middle and right, terminal
shoots showing modified flowering habit and correlation of organs after the main shoot
had been treated with triiodobenzoic acid in lanolin (5 mg./g. [middle], and 10 mg./g.).
Photographs taken 30 days after treatment.

110 LOR RENE

TABLE 5. MoLecuLar CONFIGURATION OF SOME ACTIVE AND INACTIVE BEnzorc Acips

Substances
COOH
Benzoic acid
COOH
I
2-Iodobenzoic acid
COOH
3-Iodobenzoic acid
I
COOH
4-Todobenzoic acid
I
COOH
I
2,4-Diiodobenzoic acid
I
COOH
3,5-Diiodobenzoic .acid
i I
COOH
Br
2-Bromo-3-nitrobenzoic acid
NOo
COOH
Cl

NOs 2-Chloro-5-nitrobenzoic acid

Cell
elongation

Inactive

Active

Inactive

Formative
effects

Inactive

Active

ZIMMERMAN: FORMATIVE INFLUENCES 111

TABLE 5. (Concluded)

Cell Formative
Substances elongation effects
COOH
NH»
i 2-Amino-5-chlorobenzoic acid Inactive Inactive
COOH
I
2,3,5-Triiodobenzoic acid Inactive Active
I I
COOH
Cl
2-Chloro-3,5-diiodobenzoic acid Inactive Active
I I
COOH
NHe
; 2-Amino-3,5-diiodobenzoic acid Inactive Inactive
I

that activity is dependent upon substitutions in the 2, 3, and 5 positions. None
of the mono-substituted benzoic acids were active. Also 2,3,5-triiodobenzoic
acid was more active than 3,5-diiodobenzoic acid. There are many possibilities
for substituting a given radical and combinations of various groups in the nu-
cleus of benzoic acids. Comparative activity cannot be predicted from the ap-
pearance of structural formulae. They must be synthesized and tested for com-
parative degrees of activity. That is to say, at the present time activity can be
determined only by actual biological tests.

DISCUSSION

Formative influences described under the heading of “Results” are com-
paratively new in the study of growth substances. The meaning of the word
“formative” could be extended to include local cell elongation and other short
time responses which do not involve modification of size, shape, or pattern of
organs produced by new growth. As intended in this paper, “formative” in-
volves the growth of new organs immediately following the application of the
active substance. The result is a systemic effect rather than local. It could be
compared to the effect of a systemic virus disease in contrast to a local fungus
disease. In fact the responses induced by some of the active compounds have
been mistaken for virus diseases. As the character of the responses varies with

<
by
(x)
fa
4
Oo
H

ZIMMERMAN: FORMATIVE INFLUENCES 113

different strains of virus so do they vary with the different active growth sub-
stances. A characteristic virus-like type of response is illustrated in figure 4 for
three different compounds applied to two different species of plants. The
chemically induced responses have the characteristic modifications of leaves
showing clearing of the veins, irregular shape, light and dark portions of tissue,
etc. The present results appear to lend support to the claim that virus diseases
result from natural chemical influences.

The mechanism in the plant through which these chemicals act is not well
understood. It is fairly certain that living protoplasm has many potentialities
for expressing itself and that environment determines which of these can de-
velop. The so-called “normal” characters of a plant are but a partial expression
of the range of possibilities of which the protoplasm is capable. Natural varia-
tions in the pattern of leaves of certain species of plants growing in different
environments lend support to this theory. The influences which regulate growth
must deal with undifferentiated meristems made up of uniform cells and in some
way cause them to give rise to specialized cells which in turn give rise to new
tissues and new organs of plants. Sinnott (2) is of the opinion that the cyto-
plasm plays an important role in “the construction of a pattern.” It seems
reasonable to assume that modified organs result from the influence of the
chemicals acting upon the cytoplasm rather than upon the more stable nucleus.
Each chemical constitutes a different environment and, therefore, permits dif-
ferent potentialities of the protoplasm to develop. This, at best, is only an as-
sumption but may in time help us to interpret the qualitative differences in
responses resulting from treatment with different growth substances.

The molecular configuration as a whole rather than any part of the mole-
cule appears to determine physiological activity. A slight shift in the position
of a substituted group, a change from chlorine to an amino group, or a shift in
the linkage of the chain to the nucleus may activate or inactivate a molecule.

In addition to the exact nature of the molecule, the constitution of the re-
ceptor tissue in the plant is important. First, the genetic constitution of the tis-
sue plays an important part, and second, the location in the organ and the age
of the tissue are determining factors.

Explanation of figure 3

Formative influence of 2-chloro-3,5-diiodobenzoic acid. A. Left, control; middle, term-
inal portion of tomato plant after stem had been treated with a lanolin preparation (2%)
mg./g.). Note miniature flower. Right, two abnormally large individual flowers (instead
of clusters) formed after one axillary shoot began to recover from the effects of the
chemical influence. B. Left, control; right, terminal portion of the plant which had been
given soil treatment of 2-chloro-3,5-diiodobenzoic acid (4 mg. per pot applied in 50 cc. of
water). Note abnormally small flowers on 2 clusters and recovery of axillary shoot with
abnormal flower cluster.

114 DO RIRSE WEA

Though in the same family group, tomato and potato tissues do not respond
alike to a given substance, due, perhaps to the difference in their genetic consti-
tution. Apple and lilac stem cuttings can be induced through chemical treatment
to produce adventitious roots in the spring of the year but not in autumn or
winter. Though the tissue is receptive at an early age, the capacity to respond
to the chemicals is soon lost.

B a
sa eontaaustiaas Sed
e

Fig. 4. Shoots and leaves showing formative influence of growth substances. A. Datura
stramonium: left, control; middle, sprayed with a solution of 2,4-dibromophenoxyacetic
acid (50 mg./1.) ; right, sprayed with a solution of p-aminophenoxyacetic acid (500 mg./1.).
B. Cucumber (Cucumis sativus) leaves: left, control leaf; right, 4 leaves taken from one
plant after the terminal bud had been sprayed with a solution of B-naphthoxyacetic acid
(100 mg./1.).

ZIMMERMAN: FORMATIVE INFLUENCES 115

With still other species young tissue does not respond to chemical treatment
whereas older tissue is susceptible. Many other illustrations could be given to
indicate that there are complex internal and external influences playing upon
the living protoplasm and that the sum total of these regulates the growth and
development of the plant.

SUMMARY

“Formative influence” as used in this report is defined and distinguished
from other hormone-like influences.

For the study of formative influences three groups of compounds, B-naph-
thoxy, substituted phenoxy derivatives of the lower fatty acids, and substituted
benzoic acids, stand out above all others. Substitutions in the nucleus of the
molecule may be made with halogens, amino, nitro, or methyl groups. These
may be used separately or in combination. Physiological activity depends upon
the kind, number, and position of the substituted groups. The molecular con-
figuration as a whole rather than any one part of the molecule appeared to
determine the activity.

Characteristic responses induced with these growth regulators are illus-
trated in four different figures. Attention is called to the similarities between
responses induced with synthetic growth regulators and naturally occurring
virus diseases.

Boyce THompson INSTITUTE FOR PLANT RESEARCH, INC.
YonxKers 3, N. Y.

Literature Cited

1. BoysENn-JeNsEN, P. La transmission de l’irritation phototropique dans /’Avena. Kgl.

Danske Videnskab. Selskabs. Forhandl. 1911 (1): 3-24.

Srnnott, E. W. The problem of internal differentiation in plants. Amer. Nat. 76: 253-

268. 1942.

3. ZIMMERMAN, P. W. Formative influences of growth substances on plants. Cold Spring
Harbor Symposia on Quan. Biol. 10: 152-157. 1942.

. Present status of “plant hormones.” Indus. & Eng. Chem. 35: 596-601. 1943.
(Also in Boyce Thompson Inst. Prof. Pap. 1 (35) : 307-320. 1943.)

5. ZIMMERMAN, P. W., anv A. E. Hircucocx. Formative effects induced with 6-naph-
thoxyacetic acid. Contrib. Boyce Thompson Inst. 12: 1-14. 1941.

. Qualitative differences in capacity of growth substances to induce formative

effects. Amer. Jour. Bot. 28: 14s. 1941.

. Substituted phenoxy and benzoic acid growth substances and the relation of

structure to physiological activity. Contrib. Boyce Thompson Inst. 12: 321-343. 1942.

. Flowering habit and correlation of organs modified by triiodobenzoic acid. Con-

trib. Boyce Thompson Inst. 12: 491-496. 1942.

Vo. 43 TORREYA DECEMBER 1943

Plants Need Vitamins Too*

WILLIAM J. RoBBINS

Thirty years ago when I first became interested in the nutrition of the
_ fungi the failure of a fungus to grow or grow well in a medium of known com-
position was ascribed to a variety of causes, none accounting satisfactorily for
the results. Mycologists recognized that many fungi required special media
containing some material of natural origin; and oatmeal, corn meal, potatoes,
bean pods, extract of malt, peptone, wood, dung and many other natural prod-
ucts were frequently used as such or incorporated in the material upon which
these organisms were grown. Generally speaking, an effort was made to supply
as food the material on which the organism grew in nature.

The advantage of such natural media was not understood. Some suggested
that it was because of the suitability of the minerals in the natural product, or
its favorable acidity or alkalinity, to the presence of a particular carbohydrate
or some unique source of organic nitrogen, to the special water relations
afforded by the material or to some physical property. We know now that the
growth of many fungi is conditioned by the presence in the medium of minute
traces of specific organic compounds, some of them identical with the known
vitamins ; and the presence of these growth substances in products of natural
origin frequently accounts for their advantages as culture media. This was a
possibility seriously considered by few, if any, of those concerned with the
cultivation of fungi thirty years ago. In fact, the very word vitamin was un-
known at that time; it was coined by Casimir Funk in 1912 and up to eight
years ago not a single completely convincing example of the importance of a
vitamin for a plant could be cited.

During the period from 1912-1934 the animal physiologist proceeded to
demonstrate the importance of vitamins for the growth and well-being of ani-
mals and to explore their multiplicity, functions, sources and chemistry. It
was generally agreed that plants were the sources from which animals in the
last analysis obtained their vitamins, or in other words, that plants made vita-
mins and animals used them, a fortunate circumstance for us and a sort of
philanthropic activity on the part of plants. But the possibility that vitamins
were important in the metabolism of the plant itself was regarded by the ma-
jority of plant physiologists with concealed or open scepticism. In fact, a com- |
plete and satisfactory demonstration of the importance of vitamins for plants
waited, as so frequently happens in science, on advances in another field, on
the isolation of a vitamin in chemically pure form. Crystalline thiamine (vita-

_.* Delivered at the 75th Anniversary Celebration of the Torrey Botanical Club at the
American Museum of Natural History, Wednesday evening, June 24, 1942.

116

ROBBINS: VITAMINS 117

min B,) was isolated by Jansen and Donath in 1926 and became generally
available in 1934. In that year Schopfer showed that the bread mold Phy-
comyces would not grow unless it was furnished with minute traces of this
vitamin. With this convincing demonstration as a basis and the isolation of
additional vitamins in chemically pure form our knowledge advanced rapidly.

Now we realize that plants are not so philanthropic as they once seemed.
We know they too need vitamins, but more provident than animals, most
plants make their own vitamins. Only the minority, and these chiefly the lower
plants, suffer from vitamin deficiencies; that is, they cannot develop unless
the material upon which they grow contains some of the necessary vitamins
which, of course, must come from some other kind of plant or from an animal
which has obtained them from a plant. Some bacteria, yeasts and molds need
to be supplied with vitamins. Few, if any, of the trees, vegetables, flowers and
other green plants benefit from having vitamins supplied them. To the best of
our knowledge they make all they need.

You may ask whether this means, in spite of the considerable publicly on
this subject, that supplying green plants with vitamin By or other vitamins is
not beneficial? I would answer this question in this way. The application of
vitamins to trees, flowers, vegetables and other green plants is still in an ex-
perimental stage. Some investigators have reported beneficial results on some
kinds of plants and not on others. Many have obtained negative results. We
must conclude either that the conditions under which vitamins are beneficial
to green plants are poorly understood or that their application does not bring
favorable results. Certainly the use of vitamins in horticultural practice does
not accomplish the miracles some would have us believe, and no reputable
horticulturist on the basis of the evidence now at hand would recommend their
use under normal garden and greenhouse practice.

In discussing the relation of vitamins to plans there are a good many
questions we might ask. For example, what is a vitamin, how were they dis-
covered, how do we know plants need vitamins and how many vitamins do
plants need, what plants must be supplied vitamins and what do the vitamins
do in the plant, how much of a vitamin is needed and is there a substitute for
a particular vitamin—something just as good? I can’t answer all these ques-
tions for any one vitamin, but some of them can be answered by discussing a
particular vitamin, and I have selected three, thiamine or vitamin By, py-
ridoxine or vitamin Bg and biotin or vitamin H.

Thiamine. Thiamine or vitamin B, is a white crystalline substance con-
taining carbon, hydrogen, nitrogen, oxygen and sulfur. Its empirical formula
is CyoH,gONG4S. Its structure is known and between 25 and 30 tons are now
made annually in chemical laboratories in this country. In 1935 thiamine cost
$300 per gram which is at the rate of $135,000 per pound. With the discovery

118 TORREYA

of methods of making it synthetically and the development of mass production
its price dropped to $.53 per gram or about $238 per pound. Thiamine is used
in the treatment and prevention of beri beri, of lack of appetite in children and
of various types of neuritis and in the enrichment of flour. It is perhaps the
best known and most widely advertised of all the vitamins.

The history of our acquaintance with this vitamin begins with attempts
to cure a disease common in the far east, recognized by the Chinese as early as
2697 B.C. and known as beri beri. In the 19th Century it was found that beri
beri could be- cured by controlling the diet; for example, Takaki, Surgeon
General of the Japanese Navy, about 1885. substituted meat and legumes for
part of the rice in the diet of sailors and reduced the incidence of beri beri
from between 30 and 40 percent to less than 14 percent. Takaki believed that
this was because of the increased protein furnished. In 1912 Casimir Funk, a
Polish scientist, suggested that beri beri was the result of the lack of a specific
organic substance in the food. This new dietary essential he called a vitamin.
It was found that pigeons and other animals fed on polished rice developed a
type of beri beri which could be cured by feeding rice polishings or extracts
made for them. The next step in logic was to assume that if vitamins were
really present in rice polishings and not merely in the minds of Funk and
those who believed as he did vitamins could be isolated and their chemical
nature determined. Many made the attempt. Two Dutch investigators in Java,
Jansen and Donath, succeeded in isolating a small quantity of vitamin By, in
1926, but its isolation in quantity and its synthesis in the laboratory were not
accomplished until 1934. Since 1934 many yeasts, bacteria, filamentous fungi,
and the excised roots of a number of higher plants have been found to suffer
from thiamine deficiencies. It has been found further that those plants which
grow without an external supply of thiamine make it, and the conclusion has
been reached that all living organisms need thiamine. Some make it from
simpler substances, others must be supplied with it. It is as essential and as
necessary as water or minerals or any other indispensable item in the nutrition
of an organism. Its absence means death, its presence, life.

Pyridoxine. Pyridoxine or vitamin Bg is also a white crystalline com-
pound. Its empirical formula is Cg H11O3N. It is a derivative of an ill-smelling
liquid known as pyridine. In 1939 pyridoxine cost $12.00 per gram. It can be
purchased now for $3.00 per gram or about $1350 per pound. The medical
value of pyridoxine is ill-defined. It may be of value in the treatment of certain
muscular rigidities, of paralysis agitans and perhaps of other conditions.

In 1915 Goldberger, a United States Public Health Official, recognized
that dietary deficiencies might play an important part in the development of
pellagra, a condition affecting between 400,000 and 500,000 people annually.
In 1937 Elvehjem at the University of Wisconsin demonstrated that nicotinic

ROBBINS: VITAMINS 119

acid would cure a pellagra-like condition in the dog known as “black tongue”’
and in 1938 Spies and coworkers reported that nicotinic acid was effective in
the treatment of human pellagra. During the course of Goldberger’s work he
was able to produce a syndrome in rats which he called “rat pellagra.’’ How-
ever, Gyorgy (1934 and 1935) at the Babies and Children’s Hospital in
Cleveland determined that this condition was not cured by the pellagra-pre-
venting factor, by thiamine or by vitamin Bg. It could be cured by particular
extracts of rice polishings, and he proposed that the deficiency in the food
causing this peculiar type of dermatitis was a new vitamin which he called
vitamin Bg. In 1938 vitamin Bg, later named pyridoxine, was isolated and
identified by Kuhn in Germany, Ichiba and Michi in Japan, Lepovsky in Cali-
fornia, Gyorgy in Ohio and Keresztesy and Stevens in New Jersey. It was
synthesized in 1939 by Harris and Folkers. Partial or complete deficiencies
for pyridoxine have been found for some bacteria, some yeasts and a good
many fungi. It too appears to be a vitamin needed by all living organisms.
Biotin. Biotin is a white crystalline substance which in the form of its
methyl ester has the empirical formula Cy,H;,sNe2Os35S. Its structural formula
is not yet known, and it has not been synthesized from simpler substances. It
may be obtained by a long and costly process of purification from natural prod-
ucts such as egg yolk or liver and for $10.00 you may purchase 75 micrograms
of pure biotin which is at the rate of about $62,400,000 per pound. It was first
isolated by Kogl and Tonnis of Utrecht in 1936 from the yolk of eggs and has
proved to be the most potent of all the vitamins. Kogl and Fries were able to
detect the effect on the growth of a fungus of 0.0001 of a microgram of biotin
methyl ester. Biotin is widely distributed in products of natural origin. We
have found it in such unexpected places as cow manure and cotton. In fact,
a bale of cotton contains about $1000 worth of biotin. It is made by green
plants and many bacteria, yeasts and filamentous fungi. There are, however, a
good many of the lower organisms which lack the ability to make biotin;
some cog is missing in their machinery, or it works slowly, and these organisms
grow poorly or not at all in media from which this vitamin is absent. It is
probably essential for animal growth and from recent pronouncements in vari-
ous journals may be intimately associated with the development of cancer.
The discovery of biotin has a long and interesting history. In 1860 Pasteur
published an important memoir on alcoholic fermentation in which he came to
the conclusion that yeast grew if supplied with yeast ash, ammonium salts and
a fermentable sugar. He observed that the fermentative power of yeast was in-
creased by the addition of extracts from natural products, for example, grape
juice, sugar beet juice or yeast juice but all the essentials for growth were
included, according to Pasteur, in a solution of yeast ash, ammonium salts
and glucose. However, in 1869 the famous German chemist, Justus von Liebig,

120 TORREYA

stated that yeast neither grew nor fermented sugar under the conditions de- ©
fined by Pasteur. This criticism was so keenly felt that in 1872 Pasteur de-
clared he was so sure of his results that he was prepared to perform the ©
experiment in the presence of Liebig himself. The demonstration never took ©
place, Liebig died in 1873, and the nutritional requirements of yeast as defined —
by Pasteur remained unchallenged for many years.

In 1901 Wildiers of Belgium reported that yeast would not grow under —
the conditions defined by Pasteur if the amount of yeast used in the seeding |
was small. He found that small amounts of a thermostable organic material
were necessary for the growth of yeast and gave to this chemically undefined
material the name, bios. Bios was a concentrate prepared from the yeast itself.
Wildiers suggested that Pasteur obtained his results because he had used a
large quantity of yeast for the seeding, and this large seeding had carried with
it sufficient bios to permit growth.

_ Wildiers’ proposal that minute traces of organic material in addition to
minerals, ammonium salts and sugar were necessary for yeast growth was
roughly handled by some of his contemporaries, including Fernbach (1902)
Windisch (1902) and Pringsheim (1906). Various students in Wildiers’s
laboratory supported his proposal but since no one could identify bios chemi-
cally, it remained for 20 years before the bar of science with the verdict, pro-
posed but unproven.

In 1921 MacDonald and McCollum reported that yeast would grow in a
solution of cane sugar and inorganic salts, but 2 years later Funk and Fried-
man demonstrated that ordinary cane sugar may contain a growth activator
of organic character which required for its removal three crystallizations of —
the sugar from alcohol. And so after 60 years the dispute between Pasteur and
Liebig was still unsettled. However, a decision was rapidly approaching. In
1921 Copping reported that wild yeasts would grow in a solution of minerals
and sugar while cultivated yeasts required the addition of bios for normal
growth, and in 1924 Lash Miller and Lucas of Toronto showed that there
was a difference between races of yeast in their response to bios. It seems
reasonable now to suggest that the conflict in the results obtained 60 years
before by Pasteur and Liebig may have been the result of differences in the
strains of yeast they used.

However, although the burden of evidence seemed tipping the scales in
favor of the reality of bios its chemical nature remained unknown. From 1919-
1928 various unsuccessful attempts were made to identify bios with the anti-
beri beri vitamin of Eijkman and with the coenzyme of Harden and Young
and to isolate it in crystalline form. However, Fulmer in 1923 demonstrated
that bios was not a single substance, and Lash-Miller’s laboratory in Toronto
‘separated it into two fractions, Bios I and Bios II. In 1928 Eastcott showed

ROBBINS: VITAMINS 121

that bios I was mesoinositol, a substance which more than ten years later was
found to be necessary in the diet of chicks and of rats as well as in that of
yeasts. R. J. Williams and associates separated a bios fraction which was dem-
onstrated to be thiamine and later a fraction which proved to be a new vitamin,
pantothenic acid; but a portion of bios still remained unidentified.

Kogl in Utrecht began his work in 1932 and devoted his attention to that
part of the bios complex which was adsorbed on charcoal and which he called
biotin. Four years later he announced the isolation of crystalline biotin. He had
obtained 1.1 milligrams of the crystalline material from 250 kilograms of
dried egg yolk and estimated that this amount of material originally contained
a total of 80 milligrams. On this basis it would take more than 125,000 tons
of dry egg yolk to yield 1 pound of biotin or, to put it another way, about
1,500,000 hens would have to work for a full year to produce the eggs neces-
sary to yield 1 pound of pure biotin.

But this does not end the story of biotin. About 1933 it was reported that
rats fed a diet high in raw egg white developed a peculiar and impressive skin
injury which was accompanied by emaciation and eventually terminated
fatally. This was called egg white injury. Cooked egg white did not have this
effect. It was found further that egg white injury could be cured by injections
of liver extract, and it was suggested that this was because of the presence in
the liver extract of a new vitamin which was labelled, vitamin H. In the mean-
time a group of investigators in the United States Department of Agriculture
had become interested in a factor which caused increased growth of the bac-
teria which produce nitrogen-fixing nodules on legumes. They named this
factor coenzyme R. In 1940 Gyorgy, Melville, Burk and du Vigneaud proved
that biotin, coenzyme R and vitamin H were identical.

In the same year R. J. Williams and his associates isolated from uncooked
egg white a peculiar protein, which they named avidin. Avidin it was found
combines with biotin so strongly that it renders the vitamin unavailable to
the organism. Egg white injury is, therefore, the result of a vitamin deficiency,
a deficiency of biotin and now—biotin is suspected of having an intimate rela-
tion to cancer.

Effective quantities of the vitamins. I have spoken from time to time of
effective quantities of thiamine, pyridoxine or biotin in terms of 0.01, 0.001 or
even 0.0001 of a microgram, and a microgram is one millionth of a gram. This
quantity of material cannot be seen, even with the most powerful microscope,
and it cannot be weighed, even on the most sensitive balance. It is invisible and
imponderable. If I had two dishes before me, one containing 0.001 microgram
of biotin, and the other empty you could see nothing in either dish. Yet a little
water rinsed in one dish and added to the proper medium would enable the

122 TORR Yen

proper fungus to grow while wash water from the other dish would be of no
benefit.

To the uninitiated such results border on magic, and such small quantities
are meaningless. What is 0.001 of a microgram? I will try to tell you. A tea-
spoonful of biotin weighs about 3 grams. Take one third of it and in your
imagination divide it into 1000 parts. Each part would be a milligram. Take
one milligram and divide it into 1000 parts. One of these is a microgram. It is
only necessary to think of one microgram divided into 1000 parts to obtain
0.001 of a microgram or one trillionth of a gram. Easy to do isn’t it, in your
mind’s eye?

But if such a small amount cannot be seen or weighed how can it be meas-
ured—anywhere else, that is, than in one’s imagination. This is a simple
laboratory procedure, based on the principle of dilution. If we dissolve 1 gram
of biotin in a liter of pure water it is clear that one milliliter of the solution
will contain 1 milligram of biotin. A milliliter can be readily and accurately
measured by means of a suitable pipette. If we transfer a milliliter of solution
containing a milligram of biotin to another flask of a liter of pure water and
distribute it there, then one milliliter in the second flask will contain one
thousandth part of a milligram, or one microgram. A third transfer of this
sort will yield a solution containing per milliliter 0.001 microgram or one tril-
lionth of a gram. To obtain such small quantities is easy, if one knows how.

How vitamins work. Such small quantities of the vitamins are effective
in determining the growth of an organism, like a fungus, in comparison with
the amount of some other food, such as sugar or nitrogen, that our curiosity
as to how vitamins function is sure to be aroused. It appears that they are
parts of enzyme systems, and enzymes are those substances found in the body
which make possible the chemical changes continuously occurring in a living
organism and synonymous with life itself. Much as a bit of oil speeds a huge’
machine, an enzyme makes chemical reactions go on which otherwise would
take place very slowly indeed. Sugar dissolved in sterile water will remain
unchanged indefinitely but in the presence of the proper enzyme it is broken
into its parts and yields its products. Most enzymes, perhaps all, are made up of
two parts, an enzyme protein and a coenzyme, neither of which is effective by
itself.

Some of the vitamins are known to be precursors of coenzymes. A de- _
ficiency of one of these vitamins interferes with the activity of an enzyme
system and prevents the normal metabolic changes accomplished through the
agency of that system. For example, cocarboxylase is the pyrophosphate of
thiamine. The enzyme, carboxylase, catalyzes the decarboxylation of pyruvic
acid, one of the intermediates in the metabolism of glucose; but carboxy-
_lase is only effective in the presence of its coenzyme, cocarboxylase. When

ROBBINS: VITAMINS 123

thiamine is deficient and cocarboxylase is not formed, carboxylase does not
function; and the normal utilization of sugar does not occur.

How specific are the vitamins? Vitamins are highly specific; that is,
nearly related compounds will not substitute for a particular vitamin. A small
change in the molecular structure of a vitamin reduces its effectiveness, may
eliminate its activity entirely or even change it into a harmful compound.
These results are probably because of their function as coenzymes.

What vitamins are important for plants? A dozen or more chemically
pure vitamins and similar substances are now available. Not all of these have
been demonstrated to be important for plants because usually a plant must be
discovered which is deficient for a vitamin before the need for it can be clearly
demonstrated. Nevertheless, deficiencies have been found for pantothenic acid
and para amino benzoic acid, the anti gray hair factors, for riboflavin, m-inosi-
tol, thiamine, biotin, pyridoxine and ascorbic acid. In the development of any
plant all these vitamins are probably essential, and others too, some of which
are still unidentified. Most plants, including all green plants and many of the
bacteria, yeasts and fungi, construct from sugar, minerals and a source of
nitrogen all the vitamins they require in amounts adequate for normal and
perhaps maximum development. Furnishing these plants with vitamins does
not improve their growth.

Others suffer from one or more vitamin deficiencies: that is, they do not
develop satisfactorily in a medium which lacks vitamins. Some plants have a
complete deficiency for one or more vitamins. They are unable to synthesize
any of the vitamin (or vitamins) in question, and in its absence do not grow.
This is true of Phycomyces for thiamine. Others suffer from partial defi-
ciencies; that is, they grow slowly in the absence of the particular vitamin,
but more rapidly if it is present in the medium. Apparently they are able to
make some of the vitamin, but not enough for maximum growth. Both com-
plete and partial deficiencies may be single (for one vitamin) or multiple (for
more than one vitamin). The deficiency may be absolute, or it may be condi-
tioned. By an absolute deficiency I mean that no known environmental con-
ditions enable the organism to synthesize the vitamin from the simple foods
and nutrients in a vitamin-free medium. This appears to be true of Phycomyces
in its relation to thiamine. Pythium butleri, on the other hand, suffers from a
thiamine deficiency in a concentrated mineral solution which is relieved by
diluting the solution. Its deficiency is conditioned by the medium in which it
is grown.

The synthetic ability of a plant for a particular vitamin may be complete,
incomplete, or none; that is, some plants are able to construct the vitamin
from simple food and nutrients; others are capable of making the vitamin if
supplied one or all of its intermediates; and still others are incapable of con-

124 TORE NEN

structing any portion of the vitamin. For example, Aspergillus niger has com-
plete synthetic power for thiamine; it can make this substance if supplied with
sugar and minerals including nitrates. On the other hand, Phytophthora cin-
namomt must be supplied with thiamine as such. It apparently lacks the ability
to synthesize any portion of the thiamine molecule, resembling the animal in
this respect. Between the two extremes of no synthetic power and complete
synthetic ability, there exist many types of incomplete synthetic power. For
example, Mucor ramannianus can make the pyrimidine half of the thiamine
molecule but not the thiazole portion; Sclerotium rolfsii can make the thiazole
but not the pyrimidine part; Phycomyces can combine the two intermediates
into the thiamine molecule but is incapable of making either.

The importance of studies on vitamins in relation to plants. By this time
you may be willing to admit that plants need vitamins, but you may still
question the importance of such knowledge, particularly when you remember
what I have said on the negative results of applying vitamins to green plants
in field or garden. If most plants make all the vitamins they need, why should
we study the subject?

There are many reasons. Even though vitamin B, seems at present to
exist in sufficient amounts in green plants to permit them to develop satis-
factorily without giving them more of it, there are many other vitamins, some
as yet unidentified, and we know relatively little of their relations to the
growth of green plants. If we search further, we may perhaps find a vitamin
not made by green plants in large enough quantity for their maximum devel-
opment—one which when fed to the plant wiil actually perform part of the
miracles our commercially-minded friends have told us could be accomplished
with vitamin B1; but, at present, this seems like a rather long chance.

However, it is quite necessary for us to understand the nutrition of bac-
teria, yeasts and molds, a good many of which suffer from vitamin deficiencies.
These lower plants are most important in causing disease and decay as well
as bringing benefits through their relation to fermentation, cheese making and
many similar processes. If we wish to control these lower plants—limit their
detrimental activities and encourage their beneficial properties--we must un-
derstand how they live. For example, the edible morel is one of the most de-
licious of fungi, far superior to the mushroom we buy in the markets. Yet no
one has ever cultivated it. Why? Perhaps it suffers from vitamin deficiencies
which have never been properly satisfied by the materials upon which men
have tried to grow it.

But even if further research should show that the use of vitamins on
plants is of no practical significance, the study of the relation of vitamins to
plants is important because of the light it may throw on their uses for animals.
Plants have been found to be valuable tools in determining the presence and in

ROBBINS: VITAMINS 125

estimating the quantity of various vitamins, in indicating how vitamins work
in the animal and in leading investigators to the discovery of new vitamins.
Pantothenic acid, paraamino-benzoic acid, inositol and biotin were all dis-
covered through their effects on plants before they were known to have any
influence on animals.

Entirely aside from the practical importance of using plants to increase
our knowledge of a class of substances so important for animals, or of using
the substances themselves to influence and modify the development of plants,
it should help to reéstablish self-respect in the human race at the present
moment in world history, to learn that in certain fundamental ways we re-
semble the innocent and harmless yeast plant. The same vitamins are con-
cerned in the development of yeast as in the growth of man and they probably
perform the same functions in both organisms.

And so science weaves a magic carpet of Bagdad which can carry us over
the mountains and through the jungles which once impeded and entangled the
footsteps of the seeker for knowledge. Threads from far off China, a bit of
material from Java, some from all the world are woven in its woof. There
are times when it is necessary to unravel a bit of the weaving unsuited to the
pattern, but in the end the carpet is woven, and with its aid you can scale
heights which neither Liebig nor Pasteur could surmount.

Tue New York BotTaNnicaL GARDEN
New, York, NEw York

VoL. 43 TORREYA DECEMBER 1943

Genetics, the Unifying Science in Biology*

Georce H. SHULL

One may be excused for opening a paper on a subject of this kind by several
propositions of such obvious validity that their statement is immediately recog-
nized as platitudinous :

There is no wholly unrelated fact; all truth forms a connected fabric of
inconceivably vast, indeed of infinite extent. A single observation or any num-
ber of single observations between which no connection is recognized may each
and all be true, but they do not constitute science. Science consists of a body
of knowledge which rests on recognizedly related observations. The relation-
ships between observed facts are so numerous withal, and of so many different
kinds that it is utterly impossible for any single individual to apprehend and
comprehend more than a minute fraction of all that it is possible to know.

It has been inevitable that the curiosity, that has led men to make sys-
tematic observations in order to add new facts related to those in which their
interest has been already aroused, has resulted in the sampling of many dif-
ferent parts of the network of observable phenomena and ascertainable rela-
tionships. Nature presents many different kinds of objects on which ob-
servations can be made, and among which relationships may be obviously
present or may be discovered if sufficient attention be given to them. With so
many different kinds of objects and different directions of approach there has
arisen a bewildering multiplicity of scientific disciplines, which, notwithstand-
ing their obvious overlapping and marginal merging with one another, have
tended inevitably to obscure the congruity of all facts and relationships in a
limitless universe.

Of the observations, cogitations, inductive and deductive reasoning in
prehistoric times we know nothing but there is no reason to doubt that the
human mind exercised itself in all these ways just as it does today. The history
of biological science usually starts with the marvelously comprehensive work
of Aristotle, but there must have been many pre-Aristotelians of exceptional
intellectual capacity, whose intellectual acumen and keener-than-average
powers of observation gave them high quality as individual “natural philoso-
phers,” but who, because of the lack of ready means of record and of inter-
communication, made no permanent impression on subsequent progress of
human knowledge and whose very existence can be only a matter of con-
jecture; they were the “mute, inglorious Miltons” of biological science, of
whom only a few fragmentary records, if any, remain.

* Read at the 75th Anniversary Celebration of the Torrey Botanical Club at the
Brooklyn Botanic Garden, Thursday, June 25, 1942.

126

SHULL: GENETICS 127

With the rise of recorded science came also authoritarianism,—the estab-
lishment of “schools” consisting of the students and followers of individual
observers, thinkers and teachers. Such groups of disciples did not as a rule
become independent observers, nor independent thinkers. Rather were they
simple protagonists of the theories of their leaders and disputatious opponents
of the divergent views of other leaders.

Not until the coming of the Renaissance, and the development of the
printing-press, cheapening and making more effective the process of per-
manent record and of intercommunication, could there be the accumulation of
the observations and of philosophical concepts and theories of many indi-
viduals which gradually built up the diversification of knowledge which char-
acterizes the field of science as we know it today. Along with this accumulation
of recorded fact and theory, arose the competitive spirit, the checking and re-
checking of hypotheses by new observation, the winnowing of truth from the
chaff of fallacy.

The scientific field has been enlarged by bringing new objects under ob-
servation, through exploration and importation of materials from geo-
graphical areas of ever increasing extent. Also the invention of new instru-
ments of research—the microscope, microtome, centrifuge, galvanometer, po-
tentiometer, Crooks tube, cyclotron, electron microscope,—and the discovery
of new effective chemicals, such as indole acetic acid, thiamin, colchicine, etc.,
have made possible new analyses and the perception of new relationships not
previously recognized. Similar expansion has come from the discovery of
exceptionally favorable research organisms and structures, as, for example,
the mutation phenomena, the chromosome circles, and lethal factors of
Oenothera; the regenerative capacity and tolerance of transplantation in Am-
phibians; the almost limitless genetical and cytological advantages of Dro-
sophila for studies on the relations between genes and chromosomes ; the effec-
tiveness of the coleoptiles of Avena for the recognition of growth-promoting
substances; and many others. All of these have brought about so great an
expansion of the field of biological science that ever closer specialization is re-
quired in order to make further progress. This situation has been long recog-
nized and jokingly referred to as “learning more and more about less and
less.”

So much for the expansion and diversification of biological science. As a
result the science of biology has been divided into a very large number of
separate branches, now commonly referred to as the plant sciences and the
animal sciences, plus those which relate about equally to both plants and
animals, such as general or cellular biology, ecology, and genetics.

The specialists working in each of these biological fields have found it
advantageous to organize special societies for the holding of periodical meet-

128 1 ORR YEA

ings and for the support of adequate means of record and publication of their
discoveries.

The simplest type of scientific activity is the naming and classification of
natural objects, and the first taxonomist of whom we have record was Adam,
who, according to the Biblical account, had all the plants and animals of the
Garden of Eden brought before him to receive their names. How natural that
the reawakening of human intelligence in the Renaissance should have been
characterized by the rise of taxonomy, the “mother” of all the biological
sciences! A very substantial contribution to the unification of the biological
sciences was the adoption of the binomial system of nomenclature and its very
extensive applications to both plants and animals by Carl Linné in the middle
of the 18th century.

The more philosophical phases of classification which came to recognize
natural relationships between genera, between families, and between groups
of still higher order developed more gradually and at the hands of an ever
increasing number of workers, both zoologists and botanists. On both sides
it was soon recognized that in one important corner of the taxonomic field
plant taxonomy and animal taxonomy overlap each other, so that Euglena, the
Myxomycetes alias Mycetozoa, and the Volvocineae, for example, have been
equally claimed by both plant and animal taxonomists.

Another discovery of the greatest importance for the unification of biolog-
ical science was the recognition, independently and then jointly arrived at by
Schleiden and Schwann in 1839, that both plant and animal bodies are made
up of cells and substances and structures secreted by cells. This great gen-
eralization grew rapidly in importance as refined microscopical technique
brought to light ever finer details of intra-cellular organization without finding
a single consistent difference between plant and animal cells, either in the
structures they contain or in their physiological activities.

These discoveries gave rise to the concept of biology as a single discipline,
especially through the writings and teachings of Thomas Huxley, Herbert
Spencer, John Tyndall and others. These writers emphasized the many com-
mon features of plants and animals, which made possible the stratification of
biological knowledge in fields at right angles to the taxonomic line of division
between the two Kingdoms; thus tying them together by bonds more natural
than the divisions themselves between the Kingdoms. The principles of or-
ganography, tissue-differentiations, competition and cooperation of parts,
specialization of tissues and the accompanying division of labor are equally
applicable to and derivable from plants and animals, as are all the fundamental
physiological processes, like nutrition, assimilation, growth, respiration, ex-
cretion and reproduction.

With the development of the evolution hypothesis in the first half of the last
century and its gradual acceptance by all biologists, the fact that so many

Eee eee

SHULE GH NEMICS 129

major fields of biology could bring supporting evidence, gave a still stronger
bond of unity among the several branches of biological science. Taxonomy,
comparative anatomy, embryology and paleontology were the chief sources of
this supporting evidence.

It remained to secure convincing evidence of the evolutionary processes
from actual experimentation, and here we can not over-stress the indebtedness
of the entire biological world to that great genius of simplicity, philosophical
outlook, penetrating vision and energetic persistent labor, Hugo de Vries,
whose work more than that of any other individual ushered in a new era in
biological science and philosophy. Thus was born the new experimental science
appropriately called for a time “experimental evolution,” but felicitously
christened by William Bateson in 1906, the science of “Genetics.” Inter-rela-
tionships of plant groups and of animal groups took on a new and more funda-
mental meaning when analyzed by the simple means provided by the experi-
ments of Mendel and De Vries. There came in this way a clarification of con-
cepts, and the possibility of brushing aside fallacious doctrines and their re-
placement by experimentally tested facts.

From another direction came independently another fundamental element
of genetical technology. Contemporaneous with the work of Mendel and of
De Vries was the statistical attack on problems of evolution, brilliantly con-
ceived and put into practice by Sir Francis Galton, a cousin of Charles Darwin.
This was the technique of the mathematical analysis of populations later de-
nominated “Biometry.”

Although Galton’s conclusions seemed at first to be at variance with the
discoveries of Mendel, the work of the genial and brilliant Danish plant phys-
iologist, W. Johannsen, on pure-lines and populations in beans disclosed the
nature of the discrepancy and brought complete harmony between the ob-
servations of Galton and those of Mendel and thus helped to establish biometry
as one of the fundamental biological techniques. The tool thus developed for
the handling of population problems may be considered not the least of the
contributions which genetics has made to the other sciences, most of which
tend to become more and more statistical as their stores of basic materials
grow in magnitude and diversity.

One of the most important discoveries which resulted from the experi-
ments of De Vries was the demonstration that variations, which Darwin had
taken for granted and had assumed to be more or less generally transmitted
from parents to offspring, are of two kinds. Some are completely inherited
and remain permanent elements of organization in subsequent generations
while others are non-inheritable and promptly disappear from subsequent
generations. This important differentiation of variations into inherited and
non-inherited, respectively designated “mutations” and “fiuctuations,” was
beautifully and convincingly confirmed by Johannsen, whose keen analytical

130 TORREYA

mind gave the new science of genetics its sharply accurate terminology. In-
herited variations involve permanent changes in the genotype while the im-
permanent ones involve changes in reaction of this permanent genotype under
changed environmental experiences. Only genotypic changes can have im-
mediate and direct importance for evolutionary progress, although the capacity
of a single genotype to react in different ways in response to changed environ-
ments may be of crucial importance in determining the survival of the genotype
in question in relation to its competitors in the “struggle for existence.”

Because of certain technical advantages of plants for genetical studies,
especially the facility they have for self-fertilization, Mendel’s laws were
worked out with garden peas, and all of the three nearly simultaneously pub-
lished papers of De Vries, Correns and Tschermak were based on experi-
ments with plants; but work by L. Cuenot with mice, of Bateson and his dis-
tinguished coterie of collaborators with poultry and canaries, of Long with
snails, of Castle with guinea-pigs, rats and rabbits and Davenport with poultry,
canaries and with studies of human families, quickly showed that animals as
well as plants follow identical patterns of genetical behavior.

The simplicity of the pedigree-culture methods and the fundamental im-
portance of the facts and principles to be derived from the utilization of these
methods, resulted in a very prompt participation of many investigators who
in many cases abandoned for the time being the important fields of their pre-
vious interest to become the founders of the science of Genetics as we know
it today. I have already mentioned in this connection the plant physiologist
Johannsen and the animal morphologist and comparative anatomist Bateson.
To these should be added the statisticians, Galton, Pearson and Davenport,
embryologists such as Morgan and Conklin, and cytologists like E. B. Wilson,
C. E. McClung and Calvin Bridges, to mention only a few of the more out-
standing examples.

In this way there has grown a body of knowledge of plant and animal
organization of astounding magnitude in the brief period of four decades.
There has also been demonstrated a meticulous consistency of all of the phe-
nomena which have been brought to light by these methods applied to both
plants and animals. This consistency stresses a closeness of kinship of all living

things, which hardly could have been dreamed of before the demonstration of

the genes as the elements of organization of living matter.

The genetical approach has served to bring into harmony many phenomena
of plant and animal organization and behavior which previously had had
seemingly few points in common. For example the whole field of sex relation-
ship has been greatly clarified through recognition of its basic relationship to
genetical phenomena. Mendelian heredity was soon recognized as the product
of the two critical phenomena which lie at the base of all sex, namely, the phe-
nomena of diploidization through the union of egg and sperm, and haploidiza-

a ey

SHULL: GENETICS. 131

tion brought about by meiosis, the “reduction division.” The unfortunate con-
fusion of terminology in botanical and zoological literature in relation to sex
phenomena is still only partially resolved but there can be little doubt that a
common and concordant terminology will be ultimately achieved through the
influence of genetical considerations. The confusion began when botanists
took over the sex terms which had been long applied by zoologists and by
laymen,—by the botanists themselves,—in regard to diploid animals and
applied them to the haploid generation in plants which has no counterpart in
animals.

To achieve complete harmony it is necessary only to limit the concept of
sex-homologies between plants and animals, to the diploid generation of plants,
since it is the “sporophyte” of the higher plants that manifests Mendelian
phenomena in exact agreement with those exhibited by the bodies of animals.
The situation becomes clear if we take as the starting point for a comparison
of the life-cycle of plants and animals the moment of union between egg and
sperm. This brings the diploid resting-spore of the Chlorophyceae into a posi-
tion of homology with the body of an animal, and leads to recognition of the
fact that the fundamental difference between embryophytes and animals is
the fact that in the former, the ootids (megaspores) and the spermatids (mi-
crospores) develop parthenogenetically to form respectively the female and
male gametophyte generations, whereas in animals they are converted as a
rule directly into eggs and sperms.

The closest relationship of genetics with the other biological disciplines is
that between genetics and cytology. Before the birth of genetics, cytology had
its major outlook directed toward comparative embryology. With the specific
recognition of the chromosomes as the determining mechanism of the Men-
delian phenomena, it has become obvious that cytology and genetics jointly
constitute the biology of the chromosome. Cytology represents the morphol-
ogical phase and genetics the physiological phase of the inheriting mechanism,
but the relationship is so close that it is frequently indicated by the use of the
term “cytogenetics” for this very fundamental scientific discipline.

In all other branches of biological science,—taxonomical, morphological,
physiological, sociological, psychological—the fact is of fundamental signi-
ficance that genes constitute the basic material with which the researches in
these several fields must deal. The origin and distribution of genes generally
follow a pattern of very great simplicity which must be taken into account in
laying out programs of experimentation, in analyzing the results of such ex-
periments, and in drawing tenable conclusions from them. Genetics, the
science of kinship, thus knits together, even to the most intimate details of
basic organization, the organisms with which every phase of biological science
deals, and strongly emphasizes the inherent kinship of all branches of biology.

PRINCETON UNIVERSITY
PRINCETON, NEW JERSEY

VoL. 43 TORREYA DECEMBER 1943

Criteria for the Indication of Center of Origin in Plant
Geographical Studies*

STANLEY A. CAIN

When the flora or fauna of any region is considered taxonomically or geo-
graphically, it becames apparent that it bears relationships with surrounding
regions. The taxonomist, phylogeneticist (if he be different from a taxonomist),
and the geographer are inevitably confronted by problems of origin and migra-
tion.

Forty years ago Charles C. Adams published a pioneer series of papers on
postglacial dispersal of biota in North America (Adams, 1902a, 1902b, 1905,
etc.), outstanding in their conception of process in biogeography. In one of
these papers Adams (1902a) listed 10 criteria for the determination of centers
of origin, and they were later reiterated (Adams, 1909) with further com-
ments. Insofar as I know, these criteria have never been critically analyzed,
although the concept of center of origin has been attacked by Kinsey (1936).
Rather, they have been largely accepted without question, despite the lack of
substantiating data in some cases, and have been variously and somewhat
loosely employed. It is time for an appraisal : thus it is the purpose of this paper
to review these criteria in the light of more recent contributions to the science
of plant geography. Findings in the field of genetics, in particular, and in the
study of wild populations supply reasons why certain of the criteria can not be
tacitly accepted.

The concept of center itself should be broken down into its various implica-
tions. (1) Center of origin refers only to the region in which a population or a
phyletic stock had its origin in an evolutionary sense. (2) Center of dispersal
coincides with the center of origin only for the original members of a group.
(3) Center of variation is the region where there is the largest number of bio-
types within a species, species within a section, etc. (4) Center of frequency
refers to the area with the densest population of the kind or kinds under con-
sideration. (5) Center of preservation is an area where, usually, several spe-
cies of a flora have survived a generally unfavorable change of environment.
These are the epibiotic or relic members of the flora of a region. The differences
among these centers are not always apparent in the literature.

* Read at the 75th Anniversary Celebration of the Torrey Botanical Club at the Brook-
lyn Botanic Garden, Thursday, June 25, 1942.

Contributions from the Botanical Laboratory, The University of Tennessee, N. Ser. No.
G2:

This paper has been shortened due to the space limitations of the Journal. A fuller treat-
ment of these problems may be sought in the author’s “Foundations of Plant Geography,”
to be published by Harper and Brothers.

132

CAIN: CENTER OF ORIGIN 133

The literature of plant and animal geography, taxonomy, and evolution is
replete with statements concerning the center of origin of certain species, spe-
cles groups, genera, etc. For example, Babcock and Stebbins (1937) say, “The
distribution of the genus Youngia taken as a whole is entirely consistent with
the conception that it is a natural group which had its origin in southeastern
Asia and that evolution has been accompanied by extension of the geographic
range.” On the other hand, some species, as recognized by taxonomists, may
not have had a center of origin in the sense of a restricted geographic spot
where they arose. For example, Gleason (1923) states:

. .. Probably if a complete series of specimens were at hand, showing
comprehensively the maples of the eastern states, for example, from the Plio-
cene to the present time, it would be seen that some of the earlier forms are
absolutely continuous with our present species and that the slight morphological
distinctions between them are only the result of continuous slow variations
throughout the centuries. According to this view, many modern species had no
localized origin and are not the off shoot of any parent, but represent the mass
development of a species, which, under our present taxonomic ideas, came to a
stop at the beginning of a break in our geological record of it and reappeared
as a new species at the beginning of our next experience with it.”

A different situation is emphasized by Kinsey (1936), who denies both the
usefulness and the truth of the concept of center of origin. He demonstrates
through his taxonomic work with the gall wasps that species differ by many
genic factors that have been added gradually to the population as it has mi-
grated. Some of the characters of a species have been added in one place, and
others in other places, and certain gene frequencies have increased with isola-
tion resulting from migration. Where, then, is the center of origin? I think it
would be begging the question to say that the center of origin of a species is
where the genic factor or factors causing reproductive isolation arose.

Two other situations can be mentioned in. which, in the strictest sense, there
is no single center of origin. Chromosome (genom) doubling may happen
many times in many places in a diploid population. The resulting autotetra-
ploids, which may be a good species, do not necessarily have a center of origin
other than the area of the entire progenitor diploid population. The map of
Baldwin (1941) showing the chromosome races at Galax aphylla is of interest
in this connection. Also, it is becoming increasingly apparent that many plant
species are of hybrid origin. Sometimes a swarm of diploid hybrids, segregates,
and backcrosses have attained a sufficiently distinct character and area that their
population has been given specific status, distinct from the original species. At
other times polyploid complexes develop. Stebbins (1940) says, “Dissolution
of genetic barriers and exchange of genes between genetic systems that are
completely isolated from each other in the diploid condition are made possible
by the synthesis of polyploid complexes through allopolyploidy between three,

134 A OFRERAB ING

four or more species, following the introduction of genes from all the species
concerned.” (See also Babcock and Cameron, 1934; Goodspeed and Bradley,
1942.) For example, according to the studies of Camp (1942), Vaccinium
corymbosum is a tetraploid hybrid complex that has no center of origin in the
usual sense. One contributing tetraploid was originally Ozarkian (V. arkan-
sanum), one was in the Appalachian upland (V. simulatwm), and one was of
the eastern coastal plain (V. australe).

With these qualifications concerning types of centers and with the realiza-
tion that under certain circumstances there may not be a center of origin, there
follows a consideration of the criteria proposed four decades ago by Adams.*

CRITERION 1. LOCATION OF GREATEST DIFFERENTIATION OF A TYPE

With reference to this criterion of center of origin, Adams (1902a) says, “It
is a very fundamental law that most forms of life are confined to restricted
areas and only a small number have extensive distribution. Thus, from the
center of origin there is a constant decrease, or attenuation in the number of
forms which have been able to depart far from the original home.”

This criterion is legitimate and applicable if we make two assumptions. In
the first place, the basic assumption underlying the whole thesis is that there is a
center of origin for a phyletic stock. This has already been discussed in the in-
troduction. The other assumption is that there is a time relationship in evolu-
tion, that polymorphism increases with time; and that there is an age-and-area
relationship, that with age the population of a species or other group tends to
increase and occupy a wider area. In this connection see Willis (1922, 1940)
and the numerous expert criticisms of his hypothesis. If we can accept these
assumptions, it is clear that there will tend to be more polymorphism in the
region of origin of a phyletic stock than away from this center. In such a region
there will be more forms (biotypes, subspecies, species, sections, etc.) because
of the longer time in which evolution has been occurring in the steadily increas-
ing numbers of different kinds. With time, some of the forms originating in the
central region will attain wider areas. They, in turn, may give rise to new forms
away from the center, but in the nature of the relationship, the original area
will tend to exceed any derived peripheral area in the number of kinds repre-
sented.

1] wish it understood that the evaluation of them is in no way a specific attack on
Adams’ paper, which was breaking new ground at that time, but is rather a criticism of
the present day employment of these rules without evaluation of them in the light of more
modern knowledge, and recognition of their limitations. As a matter of fact, by 1909 Adams
was careful to point out that he understood the criteria to be only “convenient classes of
evidence to which we may turn... It should be clearly emphasized that it is the conver-

gence of evidence from many criteria which must be the final test in the determination of
OASIS vee

CAIN: CENTER OF ORIGIN 135

A few quotations will illustrate this point. Payson (1922) says, “There
is much evidence for believing that Lesquerella originated at some point in
Central Texas and from this point as a center has spread over the large area
that it now occupies ... From purely theoretical standpoints also, the greatest
number of species might be expected to occur in the vicinity of the point of
origin, since there the genus would have existed for the longest period of time.”
In a recent publication on Ceanothus, Mason (1942) says, “The occurrence of
many isolated local species along the coast as against a few widespread species
of the interior would indicate that the direction of the Ceanothus migration was
from the coast to the interior.”

Another example of the use of this criterion, which also is admirably sup-
ported by phylogenetic and geological data, is the study of Gaylussacia by Camp
(1941). He says, “it becomes apparent that the genus arose in South America
for there, today, we find it as a series of interlocked species-groups still differ-
entiating out of a common plexus, only three of which have given representative
members to North America.”’ The work of Szymkiewicz (1937) indicates a
concentration of Mediterranean species of various genera, especially endemic
- species, in western Mediterranean regions. One example of this type will be
sufficient. Sirjaev (1934) has carefully mapped the distribution of the members
of the Mediterranean genus Ononis and makes the following statement con-
cerning center of origin: “Das Entstehungszentrum der Gattung (Ononis) war
wahrscheinlich auf der Iberischen Halbinsel und im nordwestlichen. Mediter-
ranen Afrika, wo jetzt noch alle Subsektionen und viele endemische und fast
alle alteren Arten sich konzentrieren, wahrend im ostlichen Teile des Medi-
terraneums keine eigene Subsektion und nur drei endemische Arten anzutreffen
sind... Die Migration aus dem Entstehungszentrum fand in verschiedenen
Epochen auf verschiedenen Wegen Statt.” The investigations of Van Steenis
(1934-1936) on isoflors (lines connecting regions of equal numbers of species
in a genus) offer another method in which a strong indication of center of
origin is obtained. Perhaps the most intensive studies of plants and their centers
ever made are those of Vavilov (1940) and his colleagues. The following quo-
tation is pertinent:

“Cultivated species as well as their closely allied wild relatives in their evo-
lution, during the course of their distribution from the primary centers of spe-
cies-formation, have been differentiated into definite ecological and geographical
groups... Primary regions are at present characterized, as a rule, by the pres-
ence of many different species (in the sense of Linnaeus). They reveal prac-
tically the entire systems of genera.”

It is necessary, however, to recognize that this criterion can not be accepted
as universal, for it only describes a tendency that, under certain conditions, is
counteracted by the operation of other factors, as is also true of age-and-area.
A few of these conditions will be described.

136 TORREYA

Requirements for the development of many species are either that the forms
are allopatric and have geographic isolation or, if sympatric, that they have
some form of genetic (internal) reproductive isolation. Regions in which there
are many closely related species are usually regions of habitat diversity, as
noted by Vavilov (1940). It is entirely possible then that a phyletic stock that
has had its origin elsewhere may, through migration, encounter a region in
which there are numerous available ecological niches that are unsaturated—
that is, in which competition pressure is low. Such a region may provide a
variety of habitats with at least partial isolation. Under these conditions a phy-
letic stock may show a “burst” of evolutionary radiation. It is apparent that such
a region of polymorphism is not necessarily indicative of the original center of
origin nor of dispersal, but is a fortuitously derived secondary center of differ-
entiation. Two more examples of this general type can be taken from Fernald’s
(1926) criticism of age-and-area. He uses the conclusions of Schonland (1924)
concerning Erica, which has nearly one thousand species in South Africa. There
is not a single known fact that indicates that the genus arose in South Africa
where there are the most endemics and the greatest diversity (species and sec-
tions). Furthermore, Willis had concluded that the number of endemics in any
genus would rise gradually to a maximum at or near the point where the genus
entered a land area, or where a genus had its center of origin. Of this corollary
of age-and-area Schonland (1924) says, “Applying this prediction to the genus
Erica in South Africa, this point would be a part of Southwest Cape Colony
west of George, where not only a large number of endemics are massed, but
where, moreover, the greatest diversity owing to formation of subgenera and
derived genera is to be found; but I fear no contradiction when I assert that it
is certainly not the place where the genus Erica entered South Africa, or where
it originated.”

Further evidence as to the care required in arriving at conclusions concern-
ing geographic problems is illustrated by Senecio. J. Small (In Willis, 1922)
localizes the evolution of the Composites through Senecio in the northern Andes
in Upper Cretaceous time, because of the present great expansion of that large
genus in the Andean region. Senecio in the mountains of tropical America is in
the most active stage of maturity, according to Greenman (1925), not because
it originated there, but because it is a region geologically young and diversified.
Small’s and Willis’ conclusion regarding Senecio rest on what Fernald (1926)
gleefully calls a “colossal geological error,’ because the present great elevation
of the Andes, where Senecio now has its magnificent development, did not oc-
cur until the close of the Tertiary (Pliocene) and the beginning of the Pleis-
tocene. From Schuchert’s (1935) recent historical geology, however, it appears
that the Cordillera Occidental and the still more western and low Cordillera de
Choco of northern South America are more ancient elevated land masses than

Te. ope? ees

CAIN: CENTER OF ORIGIN 137

the central and eastern Andes on which pre-Cenozoric plant developments
might well have occurred, and from which much of the modern Andean flora
must have been derived.

Another exception to the center of origin being where the greatest differen-
tiation of a type exists is that resulting from polyploid complexes. Polyploidy
tends to break down genetic barriers between species with a resultant produc-
tion of a large number of varieties and species (Stebbins, 1940). Examples of
such complexes include Crepis, Zauschneria, Rosa, Rubus, and sections of
Potentilla, Antennaria, and Taraxacum, and dysploidy may increase the intri-
cacy of the complex. Goodspeed and Bradley (1942) note the conclusion of
Kostoff (1938) that amphidiploids from F, hybrids may give rise to mono-
morphic species, but in other cases, if a series of segregated forms can survive,
a polymorphic species is produced. Inconstant amphidiploidy may originate a
series of adaptable forms and provide suitable material for natural selection.
In every case, according to Stebbins, the majority of the basic diploids are rel-
atively restricted in area, while most of the widespread types are polyploid. He
says, “The center of distribution of the diploid species of a polyploid complex
is naturally the center of variation of the complex as a whole .. . the diploids
tend to occupy the older, more stable habitats. This makes the study of poly-
ploid complexes very important from the standpoint of plant geography.” Such
centers of variation as are due to hybridization and polyploidy may develop at
the center of origin of a genus, but that is not necessarily the case. The Amer-
ican species of Crepis have such a center in the Pacific Northwest, but the stock
immigrated from the Asiatic center of the genus (Babcock and Stebbins, 1938).

A third type of exception to the criterion consists of such phylogenetic
stocks as have developed a center of variation at the center of origin, in the
orthodox manner, but which have suffered a decimation of the group at the
center as a result of physiographic and climatic changes. Through emigration
and extinction due to climatic and physiographic changes the variety of types
may be reduced in one region so that a secondary center comes to contain more
variety.

Hultén (1937) has also come to the conclusion that “it must... be unsafe
to assume that a plant originates in the place where it has its most numerous
relatives. In most cases such a consideration will perhaps be correct, but in
others it must be misleading.” He illustrates this point by reference to old,
widespread, arctic-montane species. “It is natural therefore that in different
parts of the area of a Linnaean species considerably differentiated races should
be found. The area has repeatedly been split up, during the glacials under the
influence of a cold climate in the north and a pluvial one in the south, and dur-
ing the interglacials under the influence of drought and heat. Each of these
agencies must have caused a selection of biotypes in its particular direction .. .

138 TORRE YEA

The idea is current that a district in which a plant shows much variation or
has many closely related species must be its original home. According to the
above point of view this would only mean that the plant has been present within
the district for a comparatively long time and has developed in different direc-
tions under the pressure of varying conditions there . . . The similarity or dis-
similarity of two types alone will hardly be able to settle discussions concerning
relationship between them.” This latter conclusion is arrived at by Hultén be-
cause of the complication resulting from “parallel selection” of biotypes by
separated but climatically similar regions.

We have seen that the location of greatest differentiation of a type may be
at the center of origin of the group and, also, that the criterion can not be un-
critically applied for a number of reasons.

CRITERION 2. LOCATION OF DOMINANCE OR GREATEST ABUNDANCE OF
INDIVIDUALS

In connection with this criterion it first is necessary to note that dominance
is a matter of the control of a community through reaction and coaction, and
abundance is only a matter of numbers of individuals. It is true that certain
forms may exert dominance through mere numbers, and that is possibly more
frequent among plants than animals, but often it is true that less abundant forms
are dominants by virtue of their life-form or strong actions.

Species that are dominants in a certain community (and there are usually
not very many such species relative to the floristic composition of the commu-
nity as a whole) usually range more widely than the area of the community.
For example, beech, sugar maple, hemlock, and yellow birch all range more
widely than the northern hardwood climax association in which they are co-
dominants. It seems to me that dominance for a species can have no meaning
except in terms of community dynamics. If, however, we consider a genus, there
may be some instances in which the regions where certain species are commu-
nity dominants or codominants are also the regions where there is a large con-
centration of species of the genus. This appears to be true for Quercus and
Hicoria in the Ozark and Cumberland regions. Even here, however, a different
interpretation is likely. These are ancient land areas in which evolution has
long been going on and the numbers of species and their dominance may be un-
related phenomena, and also unrelated to center of origin.

The center of greatest abundance of individuals, center of frequency, has
a special meaning only in connection with the distribution of the members of a
population, a subspecies, a species, etc. The assumption that the center of abun-
dance is also the center of origin for the type has to be based, it seems to me,
on an hypothesis that the species arose in the habitat where it is best capable
of abundant reproduction and establishment. This is a gratuitous assumption.

CAIN: CENTER OF ORIGIN 139

It is reasonable that, with migration from the center of origin, a species popula-
tion may encounter more favorable conditions than those that prevailed where
it arose. Hultén (1937) says concerning the “mass center’’ hypothesis, “Christ
and other authors considered that a plant is likely to have originated in a dis-
trict where its most numerous individuals are now found. Heer already opposed
this view. It is natural that if a plant at the border of its perhaps wide ori-
ginal area should find favourable conditions and multiply freely, so that
numerous individuals are developed, such a phenomenon will afford no indica-
tion of the earlier history of the species.” Such cases are apparently found in
certain weedy species of Tradescantia that have obtained wide areas and rela-
tively high abundance in the eastern grassland and agricultural areas (Ander-
son and Woodson, 1935). Also, as with criterion 1, we can conceive of climatic
deterioriation causing a reduction in numbers of individuals at the center of
origin.

Shreve (1937) has pointed out that shrubs of the Sonoran desert with
hard wood, sparse branching, and determinate growth (Cassia, Mimosa,
Acacia, Croton, Karwinskia, Caesalpima, Lysiloma, Bauhima, Acalypha, etc.)
belong to genera which are well represented in the thorn-forest, both with
respect to numbers of species and abundance of individuals. Furthermore, dis-
tributional data indicate that this type has spread from the thorn-forest into the
desert. However, Shreve (1934) has clearly shown for Larrea tridentata and
Franseria dumosa what is probably a widespread relationship—that variations
in plant size and abundance, and degree of dominance are correlated with en-
vironmental conditions, and not with the center of origin.

It is of interest to inquire further into certain characteristics of the distribu-
tion of individual plants. Gleason (1925) has studied this matter statistically
and concluded that environmental differences are not of sufficient magnitude
to affect the distribution of the species within an association, and that the num-
ber of individuals of a species, other things being equal, is an index to its adap-
tation to the environment. But what, we may ask, is the behavior of the species
outside its native association, or at the margin of its range? When the area of
a population of a new species or subspecies is expanding from its center of
origin, and when natural barriers have not yet established a boundary, there
will naturally be a centrifugal decrease of density. This would seem to be an
inevitable result of numbers and random dispersal, and to provide a case in
which the criterion is true. Let us assume, however, that a species population
has extended its area to its maximum, having met barriers of one sort or
another on all sides. Under these conditions it would seem that there would be
a tendency for a greater density of individuals to exist toward the center of
area because of a central harmony between ecological requirements and ecolog-
ical conditions. Everywhere outside of this central “typical” climatic region

140 , TORREYA

to which the species is adapted there will be, for it, a progressive deterioria-
tion of the climatic type. That is, in marginal regions of the climatic type where
it begins to grade into another climatic type, there will be fewer and fewer
suitable spots for the species. Of necessity, if this picture be true, the density
of the species will tend to decrease toward the periphery. Some interesting data
concerning the behavior of species at the margin of range have been published
by Griggs (1914) on the Sugar Grove district of southern Ohio. He says, “It
is clear . . . that in this region the species in which the individuals become
scarcer and scarcer until it fails altogether is exceptional.’ Certain species are
approximately continuous up to the margins of their range, but others are in-
creasingly discontinuous until they are characteristically disjunct, and some-
times widely so, in the peripheral portion of their areas.

In the light of these data, it would seem that the criterion of species domi-
nance and density is by no means an infallible guide to center of origin. Domi-
nance and density are frequently highly irrelevant in this respect.

CRITERION 3. LOCATION OF SYNTHETIC OR CLOSELY RELATED ForMS

From the context and through correspondence I find that by “synthetic”’ is
meant generalized or primitive forms of a phyletic group. With this half of the
criterion we can have no quarrel this far: the most primitive form or forms of
a group certainly arose somewhere, and wherever that was, there is the center
of origin of the group. To ascertain that center, after a group has had a long
history, 1s, however, another matter.

It is frequently claimed that the center of origin for a group is where the
earliest fossil forms have been found, whether or not the group is represented
there today. For example, it has been claimed that the shell family Pleuro-
ceridae had a western origin because its earliest record is from the Laramie
formation (Colorado, etc.). Adams (1915), however, concluded that the fam-
ily, and especially Jo, had a southeastern origin centering in eastern Tennessee
despite the absence of substantiating fossils.

There are two diametrically opposed views. The most widely accepted
view is that the most primitive members of a group are still to be found at or
near the center of origin of the group. This is frequently true because most of
our temperate genera date back to the Cretaceous or early Tertiary and their
primitive forms are frequently found concentrated in the old land areas. In
the United States, for example, such ancient land masses with primitive spe-_
cies (Gleason, 1923) include the Southern Appalachian center, the Cumber-
land and Ozark center, the prairie center of Nebraska, Kansas, and eastern
Colorado, the southwestern desert center, etc. In a study of Lesquerella, Payson
(1922) concluded that the center of origin of the genus was in the old land

area of central Texas where “not only are these species primitive, but in no

CAIN: CENTER OF ORIGIN 141

other locality may be found anything like an equal display of what have been
considered ancestral characteristics for purely morphological reasons .. . The
periphery in general is bounded by highly specialized members of the genus.”

The opposite view concerning the location of primitive species of a group
is that the primitive forms are to be found at the periphery of area because
they have been crowded from the center by the younger and more aggressive
members of the group. The employment of such a criterion as this depends in
part upon the validity of taxonomic criteria for the indication of primitive-
ness. Many of these criteria (as enunciated for botanists by Bessey, and others)
themselves deserve critical analysis.

One of the most skillful proponents of the view that primitive forms are
peripheral is Matthew (1939). The following quotations from “Climate and
Evolution” (pp. 10, 11, 31, 32) reveal his hypothesis which is extensively docu-
mented by vertebrate paleontology and phylogenetics, but not universally ac-
cepted.

“Whatever agencies may be assigned as the cause of evolution of a race, it
should be at first most progressive at its point of original dispersal, and it will
continue this progress at that point in response to whatever stimulus originally
caused it and spread out in successive waves of migration, each wave a stage
higher than the previous one. At any one time, therefore, the most advanced
stages should be nearest the center of dispersal (original), the most conserva-
tive stages farthest from it . . . to assume that the present habitat of the most
generalized members of a group, or the region where it is now most abundant,
is the center from which its migrations took place in former times appears to
me wholly illogical and, if applied to the higher animals as it has been to fishes
and invertebrates, it would lead to results absolutely at variance with the known
facts of the geological record . . . The successive steps in the progress must
appear first in some comparatively limited region, and from that region the new
forms must spread out, displacing the old and driving them before them into
more distant regions. Whatever be the causes of evolution, we must expect them
to act with maximum force in some one region; and so long as the evolution is
progressing steadily in one direction, we should expect them to continue to act
with maximum force in that region. This point will be the center of dispersal
of the race. At any period, the most advanced and progressive species of the
race will be those inhabiting that region; the most primitive and unprogressive
species will be those remote from this center.”

Cytogenetics is providing a body of information for several groups that
points undeniably toward the forms that are primitive in a group. One exam-
ple of this type will be sufficient. Anderson (1937) says, “In those species which
have both diploid and tetraploid races we . . . know that the tetraploids must
have originated from the diploids.” Tetraploid Tradescantia occidentalis ranges
throughout, the Great Plains and the eastern Rocky Mountains, and has a small
diploid area in central and eastern Texas. Tetraploid T. canaliculata occupies a
wide area in the Mississippi Valley, and is diploid in the same territory in

142 TORRE YeA

Texas. Also, T. hirsutiflora and T. ozarkana exhibit the same tendency. The
combination of cytology with geological history and taxonomy suggests very
strongly that the Edwards Plateau area of central Texas was the immediate
center from which the American tradescantias have developed in compara-
tively recent times.

With respect to the other point of the criterion, it can be said that closely
related forms can come to be located almost anywhere within the generic area.
The nearest relative of any form, however, will tend to be near by, at least at
first, because of the filial relationship between them. According to Kinsey
(1936), the picture of evolution is that of a simple or infrequently branching
chain. In this chain each species is a derivative of a previously existing species,
usually without extermination of the parental species.

When one looks at a large family of plants, it is apparent that it is not
everywhere equally well developed or rich. A certain tribe composing, say, 10
per cent of the family, may in one region constitute 30 or 40 per cent or more
of the family. This phenomenon is likely true for the other tribes. Such re-
gions of differentiation are likely regions of speciation or origin, except where,
for historical reasons, they are known to be regions of preservation. I can not
see, however, that closeness of relationship among species can ever be employed
as a criterion to indicate the geographic center of origin of a group without the
aid of other facts. We can only say that primitive and closely related forms may
or may not be at the center of origin.

CRITERION 4. LOCATION OF MAXIMUM SIZE OF INDIVIDUALS

In a discussion of evolution of species through climatic conditions, Allen
(1905) reiteratés some “laws” stated by him in 1882: (1) the maximum
physical development of the individual is attained where the conditions of en-
vironment are most favorable to the life of the species; (2) the largest species
of a group (genus, sub-family, or family) are found where the group to which
they severally belong reaches its highest development, or where it has what
may be termed its center of distribution.

These conclusions were reached from the observation that “in the northern
hemisphere, in nearly all types of both birds and mammals of obviously north-
ern origin, there is a gradual decrease in the general size from the north south-
ward in the representatives of a conspecific group...” Later on he says, “The
variation in size from north southward is as gradual and continuous as the
transition in climatic conditions.”

It seems to me that within these statements, employed by Adams and others,
the “cat is out of the bag.” In the first place, size is a specific character that
may not be related to environment. Size differences may be due to biotype
selection across a climatic gradient, or to phenotypic expression. Allen’s state-

CAIN: CENTER OF ORIGIN 143

ments concerning size and favorableness of environment are generally correct,
but there is no necessary relationship between size and center of origin or
center of distribution. It would seem that geographic trends in adaptive charac-
ters are usually nothing more than the clines of Huxley (1940). Allen’s state-
ments were questioned by Cockerell (1906) who said, “I found in that genus
(Hymenoxys chrysanthemoides) a case which seemed to me to exactly agree
with those postulated by Dr. Allen, except that the large form was southern,
the small one northern.” To take another case, it is a common observation
among botanists that plants on oceanic islands, such as the Azores, Canaries,
and the Galapagos, are frequently of larger stature than their relatives on the
mainlands from which they were derived. This larger size of herbs, shrubs, and
trees would seem to be related to the long growing season, rather than to any
hypothetical indication of their island origin.

I have tried to find an authentic case among plants either in favor of the
criterion or opposed to it in which the data are adequate, but have failed to do
so. The following notes are only suggestive. Prosopis, for example, attains its
largest size (height of about 50 ft.) in the Rio Grande valley, where the genus
is near its periphery. Shreve (1936) says, “It is only in the most favorable
situations that the mesquite is found as a tree. In less favorable ones it is merely
a shrub.” The genus, however, is taxonomically complicated (Benson, 1941)
and has had a long and obscure history as indicated by its split range, being in
the South American deserts as well as in Mexico and our Southwest. It is
therefore impossible to be very certain concerning the history of its area. The
Southern Appalachians are becoming famous for their large trees as the region
is better known. The largest single specimens known of Picea rubens, Tsuga
canadensis, Aesculus octandra, Tulipastrum acuminatum, and several others,
are found localized in the Great Smoky Mountains, but there is no evidence to
indicate the origin of these species in that region.

One situation in which the tendency is opposite to the criterion has been
shown by cytology. Autotetraploids, and sometimes allotetraploids, are larger
than their progenitor diploids. Furthermore, they have a strong tendency to
extend the range of the group and to occupy peripheral positions relative to
the diploids. (Anderson and Sax, 1936; Babcock and Stebbins, 1938.)

CRITERION 5. LOCATION OF GREATEST PRODUCTIVENESS AND ITS RELATIVE
STABILITY IN CROPS

From Adams’ comments, it appears that he considers productiveness to be
closely related to size and numbers, and essentially a matter of growth and re-
production. According to Adams, Hyde (1898) concluded that crop produc-
tion, whether it averages high or low, will tend to be more uniform from year

144 TORREYA

to year in the region where the crop is indigenous, and that the variability from _

year to year increases with departure from that center.* In the first place, note
that Hyde indicates that the crop production is not necessarily high at the
region of center, or where the crop is indigenous, but only that it is uniform
from year to year. This does not fit well with criteria two and four. Further-
more, it does not appear that the term “indigenous” is employed in its strict
meaning of being “native,” but in a more general meaning of being “at home”
in the sense of being well adapted. It is, of course, well known that crop produc-
tion shows the greatest stability from year to year in climatic areas to which it is
best adapted. This phenomenon appears to have nothing to do with center of
origin of the crop ( Vavilov, 1928, 1940), but is explained by weather and the
operation of limiting factors (Taylor, 1934).

CRITERION 6. CONTINUITY AND CONVERGENCE OF LINES OF DISPERSAL

When the species of a genus or higher category are distributed along natural
highways of migration, and when these highways converge on a certain area,
the distributional pattern suggests that the region of convergence of these
routes is the center of origin and dispersal. This suggestion is even stronger
when, as is usually the case, unrelated organisms show the same pattern. There
is, however, no a priori reason considering dispersal lines alone why migra-
tions need have been divergent from the apparent center rather than convergent
on it. It is usually not difficult, however, to obtain evidence (see criterion eight)
as to which direction the migrations took. Such evidence is largely obtained
from comparative morphology and relationships. Sometimes paleontological
evidence helps indicate the direction of migration. In other cases cytogenetical
analysis of the related forms reveals without doubt the direction which the
movement has taken. Migratory tracts are merely lines (however broad) of
frequent, suitable habitats, and are not necessarily one-way routes. As ex-
pressed, and by itself, the criterion is not valid.

CRITERION 7. LOCATION OF LEAST DEPENDENCE UPON A RESTRICTED HABITAT

The use of this criterion for the indication of center of origin depends upon
a species being more polymorphic at the center of origin (Criterion 1) or upon
more primitive forms having wider tolerances than derived forms. Both of
these conditions may not be true. A wide species contains a very large number
of biotypes, perhaps many thousands (Turesson, 1925, 1932; DuRietz, 1930).
Progressively from the center of origin, and especially along narrow migratory
tracts extending from the main area. there is a biotype depauperization. This
can result from partial isolation due to distance alone. A remote portion of a
population does not in practice, even if in theory, have access to the entire

? Recent investigations are summarized by Klages (1942).

CAIN: CENTER OF ORIGIN 145

stock of genes of the species as a whole. When a species is divided into geo-
graphic subspecies and ecotypes, these conditions probably apply to them also,
but less obviously. No species is completely panmictic.

On a basis of the Law of Tolerance (Good, 1931), it is concluded that each
individual organism can live only within the inherent limits of its tolerances for
the environment, and the tolerances of a species is the sum of the tolerances
of the component individuals of the species population. Now it seems to me
that this summation of Good’s can have no real meaning for an individual. No
individual can contain (inherit) all the genic variability of the population, al-
though in a panmictic population any individual might theoretically contain
any possible pair of allelomorphic genes. In many cases it is an observed fact
that morphological polymorphism decreases away from the center of area of a
species or subspecies. Although it is more difficult to demonstrate, it is reason-
able to assume that individual members of a species differ as much physiolog-
ically as they do morphologically. In fact, it seems entirely likely that adapta-
tion and ecological amplitude reside more in unseen features than in the char-
acters of the type usually employed in systematic studies. Both, of course, ul-
timately result from the genic constitution of the individuals, and may be linked.
In this connection Hiesey, Clausen and Keck (1942) say, “Within popula-
tions, hereditary variants occur, some of which may possess physiological qual-
ities that give them the potential capacity to survive in different kinds of places.
Other variations seem to have no significance for survival, representing ran-
dom differences that are not incompatible with the main requirements of exist-
ence in their population.” Just as individuals vary within a population, so may
populations show a statistical difference, which may or may not be adaptive
and favor survival. It would seem to follow, then, that when polymorphism is
greater near the center of area than at its periphery, it is entirely likely that
there will be less dependence upon a restricted habitat at the center of area. This
should not lead to the assumption that any one individual has a wider tolerance
and a lesser dependence upon a restricted habitat because it happens to live
near the center of area.

If primitive members of a group have a wider tolerance than more ad-
vanced ones, and if primitive members are more likely to be found near the
center of origin, there should be a lesser dependence upon a restricted habitat
at the center. The wide ecological tolerance that primitive species are supposed
to have is sometimes based on the paleontological evidence of large areas which
species of modern genera are known to have had in Cretaceous or Tertiary
times. This is frequently a spurious argument because many of these species
are known not to have had these wide areas synchronously, and furthermore,
little is known of ecological subdivisions of the species. Finally, there are no
physiological studies, so far as I know, which indicate that primitive species

146 . TORREYA

have unusually broad tolerances. Circumstantial evidence, on the contrary, in-
dicates that old relic species are frequently markedly restricted in area and
habitat type.

This problem has received at least one excellent consideration in paleo-
botanical literature. Ater pointing out that certain fossil floras of later Tertiary
age contain mixtures of plants from widely different habitats, Axelrod (1941)
suggests that the explanation may not be due only to overlap of floras (in
ecotonal regions, or from migratory mingling), or to the fact that Miocene
and Pliocene vegetation was “generalized” and modern forests derived by
“climatic segregation only in the late Cenozoic,” but to the ancient existence
of ecospecies. For example, Sequoia Langsdorfu (close to S. sempervirens)
was variously associated with species of boreal, warm-temperate, and temperate
type. Other modern endemics, now of restricted type, but of once wider asso-
ciation, include Lyonothamnus, Ginkgo, Glytostrobus pensilis, Picea Brew-
eriana, and Quercus tomentosa, according to Axelrod. He says, “it seems
highly probable that many Miocene and Pliocene species related to living en-
demics may represent extinct ecotypes of more widely distributed Tertiary
ecospecies.” Probable as this concept is, it still does not show that primitive
species are of wide ecological tolerance and recent ones of narrow amplitude.
The late Cenozoic was a time of climatic breakup and, for many species, bio-
type depauperization with only “senile,” relic endemics remaining; but, as
Axelrod supposes, the wide area and diversified conditions under which cer-
tain Tertiary species lived were due to the biotype (ecotype) richness of the
species as a whole. That richness represents the mature condition of a species
history. As with previous criteria, we find ourselves confronted by many “ifs.”
The above arguments concerning the region of least dependence upon a re-
stricted habitat are applicable in the determination of center of origin only
when the center of origin is also the center of variability, and when the center
of origin has not been disturbed and reduced in biotype richness.

The idea that a species is usually ubiquitous in the center of its range, oc-
curring in all kinds of places, and restricted to only the most favorable sites
at its areal limits, according to Griggs (1914), is probably attributable to
Blytt, and has been favored by Cowles. This idea includes the assumption that
the favorable climate in the central portion of the species range somehow over-
comes diverse edaphic factors, whereas at the margin of range edaphic factors
permit a spotty extension of area. I remember Cowles, when lecturing on the
dunes of Lake Michigan’s shores, saying of the cactus, “It sits on the south-
ern and western slopes, looking toward its home.” There is, of course, a large
element of truth in this generalization, as is shown by the usual disposition
of preclimax and post-climax communities in any region.

CAIN: CENTER OF ORIGIN 147

Let us turn again to the often cited polyploids. Anderson (1937) says,
“The diploid species are of limited distribution and even in those areas where
they do occur are usually restricted to one particular habitat. By contrast, the
tetraploid species and races have wide distributions and most of them have
the ability to flourish under a variety of situations.” Allopolyploids, especially,
may combine the tolerances of their diploid progenitors.

In amplifying his discussion of this criterion, Adams (1909) selects what
seems to me to be a particularly vulnerable example. He says, “Outlying colo-
nies tend to have a limited or restricted range. At the same time such colonies
are peculiarly liable to become extinct, as they are usually near the limit of
favorable conditions . . . this is true of the ‘boreal islands’ in swamps within
the glaciated portion of the continent. For example, members of the tamarack
bog association, toward their southern limit, have very restricted or local range ;
but to the north, the bog forest conditions, as it were, spread from the bogs
proper and become of extensive geographic range, as the water beetles invade
the damp mosses . . . These restricted, attenuated, or isolated colonies, depend-
ent upon special conditions, are clearly indicative that they are pioneers or
relics, which point toward the region where the range is spread out and becomes
of geographic extent.” I have italicized a portion of the above quotation to em-
phasize the fact that the areal pattern is apparently wholly dependent upon the
pattern of occurrence of suitable conditions. This is an ecological matter that of
itself denotes nothing concerning origin. Adams goes on to say that the isolated
colonies are either pioneer or relic, destroying his own thesis, it seems to me.

CRITERION 8. CoNTINUITY AND DIRECTNESS OF INDIVIDUAL VARIATIONS OR
MopIFICATIONS RADIATING FROM THE CENTER OF ORIGIN ALONG HIGHWAYS
OF DISPERSAL

This criterion, related to number six, frequently is a reliable one. With
respect to changes in character frequency (as shown by the mass-collection
techniques: Fassett, 1941) we can only conclude that there can be a gene flow
in any direction through a population. Any attenuation of the frequency of a
certain gene is presumably direct evidence of the center of origin of that gene
in the region of highest frequency. One of the most interesting cases of this
sort concerns the distribution of the recessive melanistic mutation in Cricetus
cricetus, the hamster. Timofeeff-Ressovsky (1940) says, “In the course of the
last 150 years this mutation has spread from its original center of high concen-
tration along the northern border of the species-area . . . populations with
rather high concentration of this gene are spread westward as far as the river
Dnieper.” Apparently the melanistic form is adaptive in the wood-steppe eco-
tone along the northern portion of the species area, and this is one of the few

148 ORR RIB Ne

cases in which it is definitely shown that mutations participate in the origina-
tion of geographical races.

When introgressive hybridization (Anderson and Hubricht, 1938) is dem-
onstrable and when a series of chromosome changes, such as a polyploid series,
can be shown along highways radiating from a center, it would seem that the
indication of center of origin is incontrovertible. When several characters show
a parallel and direct continuity of gradation of frequency or of modification, it
is likely that there has been active migration of the population from a center.
This is sometimes recognizable by chains of subspecies, pairs of species, etc.
Payson’s (1922) work on Lesquerella provides a good example based on com-
parative morphology. He says, “In a graphic representation of the subsectional
groups they may be shown by lines radiating from a common center. Such a
diagram could be superimposed upon a map and in nearly every case the spe-
cies at the base of each line of development would be nearer the Texas region
(center of origin) than species derived from it.” .

Once again it can be said that this criterion alone is of no significance. A
geographic series of size expressions may be due to environmental conditions
reflected in growth responses (phenotypic changes in a genotype) or it may be
due to selection operating through a region of gradually changing environment.
When morphological, phylogenetic, and geographical data are used to support
one another, the validity of the conclusions regarding direction of migration
depends upon the validity of the morphological criteria employed.

CRITERION 9. DirECTION INDICATED BY GEOGRAPHICAL AFFINITIES

This criterion is frequently valid for organisms located at stations removed
from the major area they occupy. As mentioned earlier, in any region there are
usually numerous extraneous species representing two or more different floris-
tic elements, and recording as many different migrations in the vegetational
history of the region. In this connection Grinnell and Swarth (1913) say,
“We cannot expect to derive universal laws for the behavior of species, to be
applicable uniformly in any region . . . where two faunas meet .. . Upon reflec-.
tion it is difficult to conceive of precisely the same set of delimiting factors oper-
ating upon any two species alike.” For extraneous species, it is frequently a
fairly safe assumption that they were derived from the areas where they have
their principal distribution. If a genus or family is largely characteristic of a
single formation or climatic type, and has one or a few species of different
type, it is likely that the latter migrated and evolved from the generic center.
Bromeliads have migrated away from the humid tropics and entered the deserts
of southern Mexico, and, conversely, cacti have migrated out of the desert re-
gion and established themselves as epiphytes in the tropical forest, according to
Gleason (1923). No one suspects certain rather large tropical groups as hav-

CAIN: CENTER OF ORIGIN 149

ing a temperate origin because of a few temperate representatives, as in
Diospyros, Tripsacum, and Phoradendron, but quite the contrary. The point
is well illustrated by a quotation of Merrill (1936). “When a genus is described
from material collected in a certain place and is known only from that region
for many years, we more or less automatically accept it as a group characteristic
of that region. If a representative of it is later found in another area, we are apt
to consider it as an extraneous entity there.”

Returning to our own region we can cite an example. Typical Atlantic and
Gulf coastal plain species have long been known from the Appalachian and
Cumberland uplands (Gattinger, 1901; Kearney, 1900). Sometimes these in-
land plants are rare, and stations are of small area and widely disjunct from
the coastal plain where the species are now common. Fernald (1931) has cor-
_ rectly hypothesized the origin of some of these species on the old lands that
are now part of the Cumberland plateau, and Braun (1937a, 1937b) has found
them most abundantly in the undissected portions of the now elevated peneplain.
Fernald says, “With the Tertiary uplift of the Appalachian region and its final
conversion into a vast well-drained mesophytic area . . . the Cretaceous xero-
phytes and hydrophytes which had previously occupied the ground gradually
moved out to the newly available and for them more congenial Coastal Plain
and similar habitats to the west and northwest.” In such a case as this, the prin-
cipal area is a derived one and is no indication of the center of origin. It really
is not a question of coastal plain plants in the Appalachian and Cumberland up-
lands, but of upland plants in the coastal plain, if we view the relationship his-
torically. Not all coastal plain species in the interior have had this history. In
his monographic study of the Scrophulariaceae, Pennell (1935) has detected
some forms that have migrated from the coastal plain into the Piedmont and
the Blue Ridge provinces.

The direction of dispersal and the center of origin are many times indicated
by geographical affinities, but the criterion can not be used alone, and the priici-
pal area and biographic type may be derived and the minor area relic.

CRITERION 10. DirEcTION INDICATED BY THE ANNUAL MIGRATION ROUTES,
IN Brirps

Applied to plants, this criterion would be restricted to species whose dia-
spores are bird disseminated, either epizooically or endozooically. If the migra-
tion takes place both northward and southward over the same route, as for
some species employing the Mississippi valley and others using the Appalachian
uplands, direction of plant movement is not necessarily indicated. In cases
where the northward and southward migration paths are not coincident, the
direction of movement is indicated.

150 ; TORREYA

CRITERION 11. DirEcTION INDICATED BY SEASONAL APPEARANCE

Although Adams was aware of this criterion at the time of publication of
his first list (1902a), he did not include it until later (1909). In the northern
hemisphere, vernal activity suggests bereal origin. He also thought that there
is an altitudinal as well as latitudinal relationship, i.e., that mountain forms
spreading downward should belong to the vernal aspect, and lowland forms
spreading upward should belong to the aestival aspect.

It is undoubtedly true that such relationships between origin and aspect
occur. It does not seem to me, however, that this criterion expresses any inher-
ent indication of origin. The described relationship could exist, for example,
for a form or series of forms occupying montane, subalpine, and alpine belts
(or the corresponding latitudinal zones) with the center of origin in either
terminal belt or the middle. The limitations to the spread of a form are found
in the action of the whole environment upon the physiology of the form, with
such factors as temperature, light intensity, and photoperiod operating. There-
fore, it would seem as easy and sound to conceive of a vernal form of the south
spreading northward with a change to aestival aspect, as the reverse. This fact
seems to me to illustrate perfectly the pitfialls of deductive reasoning and gen-
eralization.

CRITERION 12. THERE IS AN INCREASE OF THE NUMBER OF DOMINANT GENES
TOWARDS THE CENTERS OF ORIGIN

This criterion could only have been proposed after the development of
genetics and is appended to the older ones of Adams because of its apparent
validity. It can, I think, be attributed solely to Vavilov (1927), who said, “The
direct study of the centres of the origin of cultivated plants .. . has revealed not
only a great diversity of forms but also a prevailing accumulation of dominant
forms characterized by dominant genes in the centres. A considerable number
of plants investigated show this regularity . . . The secondary centres of the
origin of forms are, on the contrary, characterized by a diversity of chiefly re-
cessive characters.”

Several cases are discussed by Vavilov, but only one will be mentioned here
by way of illustration. The center of origin of cultivated rye and the genus
Secale to which it belongs is in Eastern Asia Minor and Transcaucasia. Here
are all the species of rye and the whole diversity of characters of the varieties ;
but also here are concentrated the dominant characters of red-eared, brown-
eared, black-eared, and marked pubescence of flowering glumes. In the second-
ary centers are such recessive characters as liguleless leaves, yellow-ears, and
glabrous glumes. Cultivated plant types in their progress from their principal
genetical centers seem to exhibit a “falling out” of the dominant genes and

CAIN: CENTER OF ORIGIN 151

“proportionally to the spread of isolation, proceeds the accumulation of reces-
sive forms.”

CRITERION 13. CENTER INDICATED BY THE CONCENTRICITY OF PROGRESSIVE
EQUIFORMAL AREAS

This criterion, developed by Hultén (1937), primarily concerns centers of
dispersal for arctic and boreal biota from refugia; but it also concerns centers
of origin when evolution as well as migration has occurred. Hultén’s thesis is
as follows: from a refugium, each species tends to spread in all available direc-
tions, but because of different tolerances and capacities for dissemination it
could not be expected that all plants would spread to the same extent or with
the same rapidity. The result is a tendency toward the development of approxi-
mately circular areas of different size around the center; but in nature the
theoretically circular form of areas is seldom attained because of various bar-
riers. There still remains, however, the chief feature of areas: those plants that
radiate from the same center have progressive equiformal areas of different size.
This criterion is obviously related to number six stated by Adams. As developed
by Hultén, however, there is a clean-cut scientific basis with the conclusion
reached through strictly inductive reasoning.

CoNCLUSION

There seems to be only one conclusion possible, and it carries implications
far beyond the scope of the present discussion of criteria of center of origin.
The sciences of geobotany (plant geography, plant ecology, plant sociology )
and geozoology carry a heavy burden of hypothesis and assumption which has
resulted from an over-employment of deductive reasoning. What is most needed
in these fields is a complete return to inductive reasoning (Raup, 1942) with
assumptions reduced to a minimum and hypotheses based upon demonstrable
facts and proposed only when necessary (Hultén, 1937). In many instances
the assumptions arising from deductive reasoning have so thoroughly permeated
the science of geography and have so long been a part of its warp and woof
that students of the field can only with difficulty distinguish fact from fiction.

Tue UNIVERSITY OF TENNESSEE
KNOXVILLE, TENNESSEE

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VoL. 43 AD OVS 18, NY AN DECEMBER 1943

Phytopathology—1867-1942*

GeEorGE M. REED

The three decades 1850-1880 are noted for fundamental discoveries in the
field of biology. In 1859 Charles Darwin published “The Origin of Species,”
a work which changed completely the viewpoint in biology. In 1865 Gregor
Mendel published the results of his experiments on inheritance in peas, an ac-
count which made no impression upon his own generation, but proved to be
the keystone of genetic investigation in the early twentieth century. Louis Pas-
teur, in 1855-1859, carried out his researches on fermentation, maintaining
that the changes which occurred in various organic substances were the result
of the activity of micro-organisms, instead of purely chemical processes in
which the observed rods were supposed to originate as by-products. In 1860-
1864 he was engaged in experiments on the problem of spontaneous genera-
tion. It was almost universally believed that the micro-organisms originated
from the decomposition of higher plants and animals. The fungi associated
with plant diseases were thought to arise from changes in the higher plants of
unknown causal origin. In 1865-1870 he carried out his classic studies on the
silkworm disease, demonstrating the microbic origin, not of one disease only,
but of two. Robert Koch, in 1876, supplied decisive evidence that anthrax of
cattle was due to a microscopic rod-shaped organism which had been associated
with this malady by Devaine and Rayer in 1850. Koch’s results were confirmed
by Pasteur in 1881, who carried out his experiments on the prevention of an-
thrax of sheep by vaccination.

L. R. and C. Tulasne, in 1861, published the first volume of their standard
work on the fungi, describing in great detail the life history and structure of
the powdery mildews. In 1863 Anton de Bary worked out the life history of a
powdery mildew, Sphaerotheca castagnei, on dandelion, describing the appear-
ance of the sex organs. De Bary’s most important work, however, was pub-
lished in 1865, when he recorded heteroecism in Puccinia graminis. Previous to
the work of Tulasne, de Bary, and others, the nature of the lower fungi was
quite misunderstood and the idea that they were the cause of various diseases
was not accepted. The demonstration of the polymorphism of the rusts, in-
volving four or five spore stages, was a great advance in our knowledge. It
was, of course, difficult for that generation to accept the view that a rust was
not only parasitic, but required at least two different hosts in order to complete
its life cycle of four or five types of spores.

* Presented at the 75th Anniversary Celebration of the Torrey Botanical Club at the .

Brooklyn Botanic Garden, Thursday, June 25, 1942.
Brooklyn Botanic Garden Contributions No. 99.

155

156 TORRE YA

The idea that a plant disease might be due to the growth of one organism
in another, resulting in the observed changes, was slow in developing. The
more common view was that the growths observed followed rather than pre-
ceded the disease, which was assumed to be due to environal conditions—the
weather, changes in temperature, moisture, and illumination, and other factors
such as the time of planting the crop, the nature of the soil, and the application
of fertilizers.

Perhaps the first disease of higher plants to be definitely connected with
the growth of a fungous parasite was bunt of wheat. Tillet (1755) provided
part of the evidence by showing that the “dust” from the bunted grains, when
applied to the seed, in some way resulted in infected wheat heads. Prévost
(1807) made a further advance by observing that the “dust” from the smut
balls resembled fungous spores and germinated in a characteristic fashion.
Kiihn (1858) also studied the germination of the smut spores and observed
the penetration of the germ tubes into the living wheat seedling. De Bary
(1863), in his early experiments on the rusts, showed by inoculation of different
spore forms that the disease followed in its characteristic symptoms. At the
time of the outbreak of the potato blight in England and Ireland beginning in
1845, Berkeley was quite insistent that the fungus observed, now known as
Phytophthora infestans, was actually the cause, although most of those with
anything to do with the disease believed that environal conditions, particularly
wet weather, were the primary factors.

Since 1867 there has been remarkable progress in working out the relation
of fungi to diseases of plants. Further, other causes of disease have been dem-
onstrated, since the bacteria are now known to produce many different types.
We also-have a whole group of diseases which are caused by a virus. So-called
“physiologic” diseases, in no way associated with a living pathogen as a
causal agent, are recognized. Many of these are due to the lack of some essen-
tial element such as boron, manganese, or some other.

It is interesting to note the parallel development in our knowledge of human
and animal diseases along with the discoveries in the plant kingdom. Koch
(1876) demonstrated that anthrax of cattle was due to a microscopic spore-
producing organism and by 1881 Pasteur had developed his vaccines for the
control of the disease. Klebs (1883) had observed the organism which causes
diphtheria, Loeffler (1884) studied the organism and obtained pure cultures,
Roux and Yersin (1888) discovered the toxin, and Behring and Kitasato
(1890) isolated the antitoxin.

The organism causing bubonic plague or “Black Death” was seen indepen-
dently by Yersin and Kitasato in 1894, and the accidental proof of its associa-
tion with the disease came in 1898. About the same time, rats and fleas were
found to be the carriers. The organism which causes tetanus was observed by

REED: PHYTOPATHOLOGY 157

Nicolaier (1884), Kitasato (1889) giving the proof of its causal connection.
One of the most striking developments was in connection with malaria, known
in various forms since ancient times. Laveran (1880) had come to the conclu-
sion that its spread was associated in some manner with mosquitoes and Ross
(1898) demonstrated conclusively that a species of Anopheles was the carrier
and, further, that the causal organism underwent cyclic changes in both mos-
quitoes and man. Soon after, the method of distribution of yellow fever was
discovered. Finlay (1881) believed that mosquitoes might be the carrier and
Reed and his associates (1900) demonstrated that the mosquito Aedes calopus
was the responsible agent. The application of these discoveries led to the eli-
mination of yellow fever as a serious disease in most parts of the world.

The story of plant pathology contains many chapters which are concerned
with disastrous diseases of economic plants. Frequently the outbreaks are due
to the introduction of susceptible hosts to new regions where an indigenous
parasite attacks them. In some cases a pathogen is carried to other parts of the
world, where it finds susceptible hosts.

The potato blight, which appeared in England and Ireland in 1845, focused
attention on this particular disease and led to great advances in plant pathology,
although the immediate results were disastrous for the people who depended
on potatoes for their food. Frequently since then potato blight has occurred in
destructive forms, and continues to be under constant investigation for methods
of control. The coffee disease, caused by Hemuleia vastatrix, appeared in Cey-
lon about 1869 and during the following years proved to be very destructive.
The final result was that the growing of coffee was given up in Ceylon, being
replaced by tea plantations, and coffee culture developed in Brazil.

The American chestnut blight was first observed in Greater New York in
1904 and the evidence is that the causal organism, Endothia parasitica, came
from the Orient on nursery stock. Since the first appearance of the disease our
native chestnut tree has been practically wiped out. The white pine blister rust
caused by Cronartium ribicola was first noted in America in 1906 on three
year old white pine seedlings imported from Germany. Previous to that, the
disease had spread widely through Europe on the American white pine, which
had been introduced. Shortly after the pathogen appeared in America it spread
far and wide on the five-needle pines and necessitated radical methods of con-
trol, which involved the attempted eradication of wild and cultivated species of
Ribes adjacent to the white pine forests. .

The rust of wheat caused by Puccinia graminis doubtless accompanied the
introduction of wheat into new regions and, wherever wheat is grown, dam-
age has been done. In the United States, 1904, 1916 and 1935 are especially
noted for the destructive outbreaks.

Since 1867 progress in plant pathology has proceeded along several lines.

158 LORE VA

1. Life history and classification of the fungous pathogens. Following
the demonstration of heteroecism in the stem rust of wheat by de Bary (1865),
the life histories of many rusts were determined. De Bary (1866) demonstrated
heteroecism in the crown rust of oats caused by Puccinia coronata, the aecial
stage occurring on species of Rhamnus. Oersted (1865) established the hereroe-
cism of Gymnosporangium sabinae. Almost every year one or more connec-
tions were established, largely by workers in Europe. Halsted (1886) and
Thaxter (1887) showed that the life cycle of Gymnosporangium juniperi-vir-
ginianae required the red cedar and the apple for its completion. Klebahn
(1888) demonstrated the connection between the white pine and species of
Ribes in the blister rust, Cronartium ribicola, and for a period of years he
devoted himself to a study of heteroecious types, by 1904 listing 178 species be-
longing to 11 rust genera. Dietel (1918) listed a total of 264 heteroecious rusts.
Arthur (1900-1921) was an active worker in growing cultures of various rusts
on different hosts in order to determine their life history, and demonstrated
that approximately 50 different North American rusts were heteroecious, in
1934 listing 153 species belonging to 14 genera in his Manual of the Rust Flora
of the United States and Canada.

Along other lines, great advances in our knowledge of the rusts have been
made. Eriksson (1894) discovered racial specialization. Blackman (1904) and
Christman (1905) described what they interpreted as a method of sexual repro-
duction at the base of the young aecial cups. It remained for Craigie (1927-
1933), in a series of papers, to demonstrate the relation of the pycnia and the
young aecia in the life cycle, thus completing the main outlines of the life his-
tory of this pathogen. There was an immediate application of these studies in
connection with the possible origination of new races of rusts.

The main facts in the life history of the bunt of wheat were established by
Tillet (1755), Prévost (1807) and Kihn (1858). A further point in the
method of distribution was brought out by Woolman and Humphrey (1924)
in which they showed that soil contamination was an important factor in our
Northwestern States.

The life history of the other smuts of cereals has also been worked out.
L. R. and C. Tulasne (1847) differentiated scme of the main types. Jensen
(1888) devised the hot water treatment for the oat and barley smuts and
distinguished two species on the latter host and Kellerman and Swingle (1890)
separated the covered smut of oats from the loose smut. Brefeld (1870-1912)
published 15 volumes recording the results of his labors on the smuts and other
fungi. Of special significance was the demonstration of the flower infection
method in the loose smut of barley and wheat by Brefeld and Falck (1905).
Zade (1924) added to our knowledge of the method of distribution of the loose
smut of oats, suggesting that to a large extent the wind-blown spores germ-

REDD PEO PAnH OLOGY 159

inated in the flowers, finally forming a resting stage, so-called “gemmae,”’ be-
neath the glumes, which later produced the infection in the young seedling.

Leveille (1851) brought out his standard work on the powdery mildews,
describing the genera which, for the most part, are accepted today. De Bary
(1863) worked out the main points in the life history, describing the sexual
organs and Harper (1895, 1896, 1905) investigated the cytology of sexual re-
production and ascospore formation. From the taxonomic standpoint, Salmon’s
monograph, published as a Memoir of the Torrey Botanical Club in 1900, was
a landmark in our knowledge of the powdery mildews.

Among the downy mildews, potato blight has been the subject of intensive
investigation wherever potatoes are grown. Studies have been concerned not
only with the pathology and the control of the organism, but also with its life
history. Berkeley, in the late 1840’s, made the first detailed studies. It remained
for Clinton (1911) to discover the oospores, Jones, Giddings, and Lutman
(1912), and Pethybridge and Murphy (1913) adding further data on the con-
ditions necessary for sexual reproduction. Gatimann (1923) brought together
the results of his detailed studies on the genus Peronospora.

Great strides have been made in the large group of the Ascomycetes and the
connection between the conidial and ascospore stages of many have been estab-
lished. L. R. and C. Tulasne (1853) described in detail the life history of

~Claviceps purpurea, which causes the ergot of rye. Aderhold (1894) and
Clinton (1901) established the connection between the common apple scab
organism and the ascocarp known as Venturia inaequalis. Norton (1902) dis-
covered the apothecia of the brown rot of stone fruits, although Schroeter
(1893) concluded that the species of fungi causing brown rot belonged in the
genus Sclerotima and Woronin (1898) showed that there were two distinct
species of this genus, S. fructigena and S. cinerea.

2. Physiologic specialization. Proper identification of hosts is basic
to an advance in the knowledge of pathogens which cause disease. Taxonomists
have been concerned largely with genera and species, while the agronomists and
horticulturists have been interested in the cultivated varieties. Students of the
parasitic fungi must necessarily be familiar with the host plants upon which
they grow since, in works dealing with their classification, the keys are largely
based upon the proper host identification, and Arthur’s recent Manual of Rusts
(1934) is a fine illustration.

One of the great advances in pathology since 1867 is the demonstration of
physiologic specialization. Schroeter (1879) called attention to this phenom-
enon in connection with certain rusts on Carex. The first important work,
however, was that of Eriksson (1894) who made an intensive study of Puccinia
graminis from the cultural standpoint. On the basis of his experiments, he rec-
ognized 6 formae speciales—Avenae, Secalis, Tritici, Airae, Agrostidis, and

160 A OPRIRIEAV er.

Poae. Another step was taken in 1917, when Stakman and Piemeisel found that
P. gramuinis tritici consisted of at least more than one specialized race or physio-
logic form. By 1922, 37 specialized races of this pathogen were known and by
1934 not less than 127 had been isolated, and now the number is about 160.
Similar specialization has been found in other groups of grass rusts. In crown
rust of oats Murphy (1933) listed 33 races and Johnston et al (1942) brought
together the data for Puccimia rubigo-vera tritici, recording 129 races known in
various parts of the world. Most rusts which occur on several species of grasses,
particularly if they belong to different genera, show the phenomenon of special-
ization. It is interesting, however, that Puccinia subnitens does not, the spores
from the uredial and telial host being able to infect aecial hosts belonging to 15
genera, distributed among 6 different families.

Specialized races of the powdery mildews were first recorded by Marchal
(1902) when seven were differentiated on the basis of cultural experiments—
Avenae, Agropyrae, Bromi, Hordei, Poae, Secalis, and Tritici, all being limited
to one or more species of a single genus. Salmon (1903) and Reed (1906-
1916) extended the evidence for specialization within this mildew. A further
step was taken by Mains and Dietz (1930) when they showed that Erysiphe
graminis hordei consisted of at least 5 distinct races, and Mains (1933) found
2 races of E. graminis tritici.

The first evidence of specialization in the smuts was recorded by Zillig
(1921) in Ustilago violacea. Faris (1924) demonstrated the occurrence of 5
physiologic races in the covered smut of barley, U. hordei, and Reed (1924)
demonstrated races in both loose and covered smuts of oats. At the present time
30 specialized races of loose smut and 14 of covered smut are known. Faris
(1924) demonstrated specialization in the bunt of wheat, his data being ex-
tended by Reed (1927, 1928) when 5 races of Tilletia levis and 6 of T. tritici
were differentiated. Rodenhiser and Holton (1937) listed 8 physiologic races
of T. levis and 11 of T. tritici. Such specialization has also been found in
Sphacelotheca sorghi, Sorosporium reilianum, Ustilago tritici, and U. zeae.

Physiologic specialization is an essentially universal phenomenon among
the pathogenic fungi. In any case where a morphological species of a fungus
occurs on several hosts, it is almost certain that strains or races exist which
are limited in their capacity for producing infection.

Ward (1903), in his study of the brome rust, Puccinia disper Sa, raised ae
question whether “bridging hosts” existed, publishing data which he regarded
as evidence that a particular race of brome rust might be grown on a specific
host and then be capable of infecting other brome grasses which originally it
was not able’ to do. Salmon (1904) published similar data for the powdery
mildew on the brome grasses. For many years no clear-cut confirmation of
these conclusions was available. The general idea, however, was held in con-

REED: PHYTOPATHOLOGY 161

nection with the rusts that the aecial host might be a meeting place for different
races, resulting in a changed capacity for infection in the uredial stage. It is
now known that on the aecial host hybridization of the races of the pathogen
may take place, and thus new races arising might differ greatly in their capacity
for infection.

Reddick and Mills (1938), in connection with the potato blight organism,
have suggested that when it is grown on partially resistant hosts, it may ac-
quire an ability to infect a wider range of varieties, thus bringing to the fore
again the question of bridging hosts.

3. Environal factors. Before 1867 the view was that environal factors
were the principal cause of plant diseases and, as a corollary of this, the fruit-
ing bodies of the fungi which appeared upon the plant followed the disease.
Epiphytotics, such as the potato blight in the 1840’s were largely attributed to
the weather.

We now recognize the very great importance of environal factors as pre-
disposing the appearance of a diseased condition ; in fact, three different things
are necessary: (1) a susceptible host, (2) a causal agent such as a fungus,
bacterium, or virus, and (3) environal factors that are favorable for the es-
tablishment of the relation between the two. We must emphasize the interrela-
tions of environal factors, including soil temperature, moisture, and reaction,
since it is impossible to find a fixed optimum for any one, regardless of the
possible associated variables.

While we know that the real cause of many diseases is due to specific or-
ganisms, we also know that particularly disastrous epiphytotics occur only
under peculiar environal relations. Jones, Giddings, and Lutman (1912)
worked out the relation of weather conditions to the development of potato
blight. The prevalence of wheat bunt depends upon low soil temperature at the
time of seeding. Oat smuts are not as destructive, ordinarily, in the Eastern
United States as in the Western. !

Intensive studies on the relation of environal factors to plant diseases have
been made. The relation of temperature and moisture to the infection of wheat
by the two species of Tilletia was made by Hecke (1909), Heuser (1922),
Munerati (1922), Hungerford (1922), and Faris (1924). Faris (1924) stud-
ied the temperature and moisture relations for infection of barley by the
covered smut, Bartholomew and Seymour (1923) for the loose smut of oats,
and Reed and Faris (1924) for the covered smut of oats and the loose and
covered smuts of sorghum.

On the establishment of the Department of Plant Pathology at the Univer-
sity of Wisconsin, Professor L. R. Jones and his students conducted exten-
sive studies over a period of years, with elaborate equipment, on the influence
of environal factors on the development of many plant diseases. While empha-

162 WO RRE YA

sis was laid on temperature, other factors such as moisture and soil conditions
were determined in the case of cabbage yellows, flax wilt, tomato wilt, tobacco
root rot, stem canker of potato, and other diseases. An interesting result was
observed in seedling blight of cereals caused by Giberella saubinetii, an organ-
ism causing the disease in both corn and wheat. In corn, severe infection occurs
at 16° C. and only slight infection at 24°, while in wheat the temperature rela-
tions are reversed. Jones, Johnson, and Dickson (1926) have summarized the
investigations.

Seasonal development influences the reaction of many plants to a particular
disease. Waterhouse (1929) found that barley hybrids gave different results
in winter and summer months, when inoculated with Puccinia anomala. Some
families in winter gave a normal ratio of 3 resistant to 1 susceptible, while in
summer the progenies failed to show the expected segregation. Harrington
(1931) found that a series of progenies of a cross of Marquillo X Marquis
showed susceptibility as dominant with a race of P. gramunis tritici at a high
temperature, while at a low temperature resistance was dominant. Mains
(1934) found that hybrids between Michigan Amber and Chinese wheat were
difficult to classify in their reaction to a race of Erysiphe gramunis tritici when
grown in the spring, while it was easy to group the hybrid lines when grown
in the winter. One parent, Michigan Amber, was resistant in the winter and
more or less susceptible in the spring. Gordon (1930, 1933) found that some
oat varieties showed no significant differences in their reaction to certain
physiologic races of Puccinia graminis avenae when grown at four different
temperatures from 57.4° to 75.4° F. The Joanette variety, however, was
very resistant to some other races at low temperatures and susceptible at high.
Peturson (1930) found that Red Rustproof oats was resistant to a race of P.
coronata avenae at 57° and susceptible at 70 and 77°. Four other varieties were
fully susceptible at all three temperatures, while a fifth variety was resistant.
Another aspect of the problem was brought out by the work of Goulden, Newton
and Brown (1930). Some wheat varieties showed no essential differences in
reaction to particular physiologic races of P. gramunis tritici in the seedling and
in the mature plant stage. Other varieties, however, differed markedly in re-
sistance in the two stages of plant growth. These results have been confirmed
by other investigators.

4. Diseases caused by bacteria and other organisms. In addition to
the diseases of plants caused by fungi, it is now known that many important
diseases of plants are caused by bacteria and other organisms.

Let us recall the fact that Koch (1876) demonstrated conclusively that
anthrax of cattle was caused by bacteria. In the period 1878-1883 Burrill car-
ried out his studies which showed the relation of fire blight of pears to particular
bacteria. Then followed in rapid succession other demonstrations of the rela-

REED: PHYTOPATHOLOGY 163

tion of bacteria to plant diseases—Wakker (1883-1889), yellow disease of
hyacinths; Smith (1897) and Russell and Harding (1898) black rot of cab-
bage; Stewart (1897) bacterial wilt of sweet corn; Smith and Townsend
(1907) and later publications by Smith and others on crown gall. These, and
such other diseases as blight of beans, citrus canker, soft rot, cucurbit wilt, black
leg or black rot of potato, red-stripe disease of sugar cane, and wildfire of to-
bacco, have all been associated with bacteria. Smith (1905, 1911, 1914) pub-
lished three large volumes dealing extensively with the bacterial diseases and in
1920 published his summary. Elliott (1930) listed 177 species of bacterial plant
pathogens—13 caused by Aplanobacter, 53 by Bacillus, and 111 by Bacterium.

It is also interesting to recall the controversy between Dr. Alfred Fischer
and Dr. Erwin F. Smith in 1899. The former maintained that bacteria did not
cause disease in plants, while Smith affirmed their causal connection.

Other organisms have also been associated with plant disease. Club root
of cabbage, caused by Plasmodiophora brassicae, has been studied by Woronin
(1878), Lutman (1913), Kunkel (1918), and others. Root knot or root gall,
caused by nematodes, was first observed by Berkeley (1855). Greef (1872)
described the nematodes, Frank (1885) and Atkinson (1889) gave further
details on the disease and the causal organism. The nematode disease of wheat
was found by Johnson (1909) in California and by Fromme (1917) in Vir-
ginia. Byers (1918-1920) has made detailed studies.

5. Virus diseases. A separate chapter in plant pathology deals with the
virus diseases of plants. The first scientific studies were concerned with the
tobacco mosaic, which has continued to be a favorable subject of many investi-
gators. Mayer (1886) discovered the infectious nature of the juice of mosaic
tobacco plants, [vanowski (1892) discovered that the infectious principle could
pass through a Chamberland filter, which held back bacteria, and Beiyerinck
(1898) extended the work, introducing the term “contagium vivum fluidum.”
Many plant diseases are caused by a filterable virus, among them aster yellows,
curly top of beet, sugar cane F'1ji disease, peach yellows, stunt disease of rice,
mosaics of sugar cane, cucumber, hop, lily, and potato. We may note in passing
that Loeffler and Frosch (1898) established the first causal connection of a
virus to a disease of animals, the foot and mouth disease of cattle.

Studies have been made on the methods of transmission of the filterable
viruses, being distributed by grafting, budding, and on the seed, as in the case
of the legume mosaic. A most interesting development is the discovery of insect
vectors. Takami (1901) found that the stunt disease of rice which was often
destructive in Japan, sometimes resulting in crop failures involving famines,
was caused by the feeding of the leaf hopper, although the actual virus was not
discovered until 1908-1909. Aphids and leaf hoppers are very common vectors.
Usually, there is a high degree of specialization in the carrier, a specific insect

164 LORREYA

being responsible for a particular disease. Remarkable progress has been made
in the study of the nature of the viruses. Duggar, Kunkel, Smith, Stanley, and
many others have made important contributions.

6. Disease resistance. From the earliest times it was observed that
species and varieties varied in their susceptibility to disease, and the resis-
tant ones were selected in order to minimize loss. In recent years great progress
in the selection of these has been made and programs have been developed in
the field of plant breeding for combining the resistant quality with other desir-
able characters. Success is dependent upon the close cooperation of the plant
breeder and the pathologist.

Orton (1899 and later) stressed the value of types of watermelons resis-
tant to the wilt disease and by 1913 had developed commercial varieties. Norton
(1910) obtained varieties of asparagus resistant to the rust. Jones and Gilman
(1915) began their work on cabbage resistant to yellows. Edgerton (1918)
and Pritchard (1922) have developed wilt-resistant tomatoes. Jagger and Scott
(1937) obtained cantaloupe varieties resistant to the powdery mildew.

Finding resistant stock is the first step in any breeding work. The species
or varieties may be brought in from other countries and used in the program.
Wild potatoes have been sought in Mexico and Peru, and melons from India
have proved useful. Graves is finding chestnuts from the Orient useful in devel-
oping hybrids of our native chestnut which are resistant to blight. Barley, oat,
and wheat varieties have been carried from one part of the world to another
and serve as basic stock in breeding programs.

In most groups of economic plants, studies on varietal resistance have been
made, for example: Reed, Griffiths and Briggs (1925) on the resistance of oat
varieties to both loose and covered smuts, Reed and Melchers (1925) on the
resistance of sorghum varieties to the covered smut, and Tisdale et al. (1923)
on the resistance of varieties of wheat to the flag smut, and (in 1925) to bunt.
At the Institut fiir Pflanzenbau und Pflanzenzuchtung, Halle-Saale, students
of Director Th. Roemer have made similar studies of several of the cereal smuts.

Rieman (1939) stated that about 80 resistant varieties of vegetable crops
had been developed and at least 20 of these were recognized by the trade, in-
cluding asparagus resistant to rust, snap beans to mosaic, cabbage to yellows,
corn to Stewart’s bacterial disease, lettuce to brown blight and powdery mil-
dew, peas and tomatoes to fusarium wilt. Coons (1937) estimated that about
one-quarter of the acreage devoted to 17 important crops in the United States
was planted to disease-resistant varieties.

Breeding for disease resistance is a difficult and time-consuming procedure
and there are many hazards by the way. Frequently new physiologic races of
the pathogen appear. This is well illustrated in potato breeding for blight re-
sistance. The first attempts to obtain resistant varieties were made in the late

INAADs Ieee WiakOlLeOxrere 165

1840’s and ever since efforts have been continued to secure resistant varieties.
In a few cases promising results were secured, especially when a new breed-
ing stock was obtained from Mexico or Peru. Since 1918, Reddick in the United
States, Salaman in England, Muller and Schick in Germany, and workers in
Russia, have developed blight-resistant potato breeding programs. However,
the discovery of specialized races of the pathogen in 1933 by Miller and by
Miss O’Conner and Peterson (1933) have made the program more difficult.
Another example of the difficulties in the successful development of resistant
varieties is found in breeding oats for smut resistance. The variety Victoria was
imported from Uruguay by the United States Department of Agriculture in
1927. After its introduction it proved to be resistant to all races of loose and
covered smut known at that time. It was crossed with other varieties and by
1940 many valuable selections had been obtained which combined smut resis-
tance with other desirable qualities. The discovery of a new race of smut in
1941, which attacks Victoria and most of the selections derived from its crosses,
necessitates a new breeding program.

The genetics of disease resistance has been investigated by many workers.
Biffen (1904) early published data on the yellow rust of wheat, Puccinia
glumarum, which indicated that the inheritance of resistance followed the Men-
delian laws. The rusts have been suitable for such studies, since the results from
an experiment may be secured in seven to ten days. However, environal factors
must be carefully considered. Many hybrids have been studied by Hayes et al.
(1920), Harrington and Aamodt (1923), Clark and Ausemus (1928), Goulden
et al. (1928), McFadden (1930), as well as other investigators. Sometimes
the results have indicated a simple relation, while in others the genetic situation
is quite complex.

The smuts of cereals have been favorable subjects for the study of the inheri-
tance of disease resistance. One of the difficulties, however, is the long period
of time required for securing the data, and another is the great importance of
the control of environal factors at the time of infection. Gaines (1923) obtained
a complicated situation in his studies of the genetics of bunt resistance. Briggs
(1926 and after) secured quite clear-cut results which usually indicated mono-
hybrid ratios. He reported, however, the occurence of several factors for resis-
tance found in different varieties. Crosses between resistant and susceptible
varieties of oats have been studied with reference to their resistance to loose
and covered smuts, beginning with Wakabayashi (1921), Gaines (1925), and
Reed (1925). Many different hybrids have been studied by workers, and the
results sometimes indicate clearly a single factor difference, while in other
crosses two, three, or even more factors are required to explain the data. Mains
(1934) studied the resistance to powdery mildew of wheat hybrids and Briggs
(1935 and later) carried out a series of experiments with different hybrids of

166 TORREYA

barley, sometimes obtaining simple relations, but identifying several distinct
factors for resistance to a specific race of the powdery mildew.

7. Disease control. Viewed by the practical man, the control of disease
is the primary consideration, and the emphasis is placed on securing adequate
methods for avoiding the losses due to the destructive diseases. The selection
of resistant varieties is one method of procedure, but many others have been
employed. The prevention of disease, rather than an attempted cure of infected
plants, is recognized as of first importance. Ward (1882), in connection with
the coffee disease, clearly emphasized the idea of preventive treatment. It is
essential that the toxic material be applied so that it is on the leaves when the
spores of the pathogen are germinating. Whatever material is used, it must be
applied at the right time.

In a few cases curative measures are successful. In loose smut of wheat
and barley, the invasion of the parasite occurs just after the period of polliniza-
tion and as the grain ripens the fungus passes into a dormant condition, and
may be killed by the hot water treatment. There are a few other illustrations,
particularly in the case of virus diseases, as discovered by Kunkel.

Previous to 1867 there were two diseases of plants which were more or
less effectively controlled by chemical substances. One was the powdery mildew
of the grape by the use of sulphur, discovered by Tucker (1847), and the other
the bunt of wheat by a method of seed treatment with salts of copper, as worked
out by Prevost, Kiithn, and others. Since 1867 great strides have been made
in the control of diseases by chemical means. Many sprays and dusts have been
utilized, one of the most important being Bordeaux mixture, discovered by
Millardet in 1882, which was effective against the downy mildew of the grape.
This spray, with modifications, is still one of the standard materials in the
control of many diseases. Lime-sulphur was accidentally discovered in 1885
as an effective control of the peach leaf curl, Pierce (1900) giving the history
of its use. Scott (1908) reported experiments on the value of self-boiled lime-
sulphur, which was effective in the control of peach scab and brown rot, and
was successfully used to control apple scab in 1910. Great emphasis has been
placed upon the use of dusts instead of sprays in the control of fruit diseases.
Whetzel and his associates have been active in the development of suitable
dusts.

Copper, mercury, and sulphur remain, at the present time, the vekneiael
materials for the chemical control of disease. However, great advances have
been made in the use of these elements in new types of compounds and in the
physical make-up of the dust or the spray. Investigations have been carried out
on the proper methods of applying the material, the discovery of suitable
spreaders and stickers, and methods of control involving the combination of
insecticides and fungicides. Important changes have occurred in developing

= Ree

REED PHY TOPATHOLOGY 167

suitable spraying and dusting machinery, and elaborate schedules for applica-
tions for the control of various diseases and insect pests have been worked out.

Great advance has been made along the line of seed treatments. Formal-
dehyde was first successfully used by Bolley (1897) for the control of oat smut,
and Haskell (1917) devised the spray method, thus solving the problem of the
wet grain. Copper carbonate dust was introduced for the control of bunt of
wheat by Darnell and Smith (1915) in Australia and Mackie and Briggs used
this material successfully in the United States in 1920. Riehm (1913) dis-
covered the value of organic mercury compounds, as chlorphenol mercury, in
the control of smut diseases. Important advances in the use of the organic
mercurials have been made, utilizing such substances as uspulun, germisan,
chlorophal, and semesan.

The application of heat has proved successful in the case of some diseases.
Jensen (1882) partially controlled the potato blight by heating the tubers. In
1888 he applied the hot water method to the seed of oats and barley for the
prevention of smut. The hot water method was improved by Appel and Riehm
(1911) and by the pathologists in the United States Department of Agricul-
ture since 1920. Kunkel (1936) found that heat treatment is effective in the
control of peach yellows, diseased plants recovering after being held for some
time at 35° C. The yellows of periwinkle disappeared if infected plants were
held 38°-42° C. for two weeks (1941).

8. Research and teaching. With rare exceptions, previous to 1867
botany was not recognized as an important subject for research or instruction
in colleges and universities. Little attention was paid to pathology, most of the
work being done in Europe. Since 1867, however, research and teaching have
greatly expanded, not only in Europe but also in the United States. Thomas
Taylor was appointed microscopist in 1871 in the Department of Agriculture
and in his first report published an illustrated article on the diseases of grape,
pear, and peach trees and lilacs. In 1886 a Section of Vegetable Pathology with
Frank Lamson-Scribner as Chief was organized in the Division of Botany, and
the first bulletin was on the fungous diseases of the grape vine. E. F. Smith,
an assistant in the Division, started his investigations on peach yellows, the first
bulletin on this disease appearing in 1891. Farlow (1874) began his investiga-
tions and teachings along pathological lines. Burrill (1878) began his studies
on pear blight.

In 1888 B. T. Galloway was appointed Chief of the Division of Vegetable
Physiology and Pathology, heading the Bureau of Plant Industry when it was
established in 1901. Further reorganization of the botanical and pathological
work of the Department has taken place, but diseases of plants continue to oc-
cupy the time of many investigators. The importance of pathology is empha-
sized by the organization of the Division of Cereal Crops and Diseases, Division

168 TORREYA

of Fruit and Vegetable Crops and Diseases, and the other Divisions of the
Bureau of Plant Industry.

The Rockefeller Institute for Medical Research in 1932 established at
Princeton, New Jersey, a laboratory for research in plant pathology, an institu-
tion largely devoting its attention to the virus diseases of plants.

In the State Universities, Agricultural Colleges, and Experiment Stations,
the study of plant diseases has been given increased attention. Before 1900, the
botanists of the institutions may have carried on investigations on some diseases
of plants. Later, men were appointed to devote their entire time to pathology.
No State, however, had a pathologist until after 1900, although fine pathological
work was done by Burrill, Arthur, Jones, and others. The first separate De-
partment of Pathology was organized at Cornell in 1907 under Professor H. H.
Whetzel. In 1909 Professor L. R. Jones headed the Department of Plant Path-
ology at the University of Wisconsin. In California Dr. R. E. Smith in 1903
was appointed Assistant Professor of Plant Pathology in the Department of
Botany, and in 1907 Dr. E. M. Freeman received the title of Assistant Profes-
sor of Botany and Pathology at the University of Minnesota. Pathology, in
most institutions, is a part of the Department of Botany, although in a few it is
separated.

There has been a great increase in the facilities for the encouragement and
publication of research. The American Phytopathological Society was founded
in 1908 with about 200 charter members, the enrollment in 1941 consisting of
1120 members.

Most botanical journals publish papers on plant pathology. A few, however,
are devoted largely to this phase of botany: Zeitschrift fur Pflanzenkrankheiten
(1891) edited by Dr. Paul Sorauer; Phytopathology (1911) first edited by
L. R. Jones; Société de Pathologie Végétale de France (1914) ; Review of
Applied Mycology (1922) edited by E. J. Butler; Phytopathologische Zeit-
schrift (1929) edited by E. Schaffnit.

The Bureau of Plant Industry, United States Department of Agriculture,
from 1901-1913 published 285 Bulletins, as well as Circulars, many of which
were devoted to pathological subjects. The Journal of Agricultural Research
succeeded the Bulletins in 1913, and has published many papers along patho-
logical lines. In addition, the Department still continues to issue Technical
Bulletins in pathology, as well as in related botanical and agricultural fields.
The Agricultural Colleges and Experiment Stations have issued many Circu-
lars, Bulletins, and Memoirs, on plant diseases.

Previous to 1867 there were very few textbooks dealing with pathology.
Among the earlier were those of Unger (1833) ; Weigmann (1839) ; Meyen
(1841) ; Berkeley (1854-1857) ; and Kithn (1858). Sorauer published the
first edition of his Handbuch der Pflanzenkrankheiten in 1874, consisting of a

REED: PHYTOPATHOLOGY 169

single volume. In 1933 the first volume of the sixth edition of the greatly ex-
panded work appeared. Hartig (1882) published his text on tree diseases.
- Kirchner (1890), von Tubeuf (1895), and Frank (1896) wrote general texts.
Since 1900 many texts have been published, among the first being Duggar’s
Fungous Diseases of Plants (1909). Some of the texts cover the general field,
while others are limited, dealing either with diseases of fruit trees, vegetables,
cereals, ornamental plants, or trees.

One of the most important developments in the advancement of plant path-
ology and the control of plant diseases was the passage of legislation. Great
Britain (1877) passed its Destructive Insects and Pests Act against the Colo-
rado potato beetle and, in 1907, against all insect pests, the first ruling being
applied against American gooseberry mildew and the wart disease of potato.
The United States Department of Agriculture (1912) established a Federal
Horticultural Board and issued the Quarantine Act. The first orders were
against white pine blister rust and the wart disease of potato.

BrooKLyn Boranic GARDEN
Brooktyn, NEw York

At the meeting at the Brooklyn Botanic Garden on Thursday, June 25, a
fourth paper was presented by Dr. A. F. Blakeslee on “Technical Applications
of Genetics in Plant Breeding in 75 Years.” Unfortunately this paper is not
available for publication.

VoL. 43 TORREYA DECEMBER 1943

The Field Trip to the New Jersey Coast and Pine Barrens
Friday and Saturday, June 26-27, 1942

E. J. ALEXANDER AND H. K. SvENSON

As in the case with most field trips the participants came from many direc-
tions by train and automobile, to join at Point Pleasant. The early departure
from New York had left most of the group without breakfast so that an hour
or so was squandered in the various cafés of the village, but Doctor Chrysler
finally rounded up a party and we proceeded along the railroad track and
road to the south of Point Pleasant. We had gone perhaps a quarter of a
mile, noticing the large trees of Quercus phellos on the roadside, when we
were pulled up into a meadow on the east side of the track. This meadow
had a good many of the interesting plants to be found along the seacoast above
tide-level, such as the two milkweeds, Asclepias rubra and A. lanceolata, the
latter species apparently reaching its northern limit at this point. There was
much interest in the yellow flowers of Oenothera (Kneiffia), but all the varia-
tions seemed to resolve themselves into one species, O. longipedicellata. The
meadow also had a good deal of Aletris farinosa, the white spikes being es-
pecially conspicuous at this time of year, and some scattered plants of Polygala
lutea, a species which is more at home in the pine barrens. A large colony of
Viola Brittoniana was found here, the plants in full seed. This is an attractive
cut-leaved inhabitant of acid coastal soils, rather rare and localized in its occur-
rence, so that a future trip was planned for the following spring to see the
colony in flower.

Making a short turn toward the ocean we came to one of the lagoon-like
ponds bordered by a wealth of interesting aquatics. Creeping along the shore
were Myriophyllum tenellum in great abundance and also the more common
M. humile ; along with a carpet of the small yellow Utricularia gibba, Gratiola
aurea, Eriocaulon septangulare, Hydrocotyle umbellata, and Elatine ameri-
cana. At the margin of the pond were several specimens of Ranunculus scelera-
tus, an interesting species with exceedingly acrid juice and rare in the New
York region. Farther out in the water, to be reached only by deep wading, was
a growth of Potamogeton pectinatus, a species generally of limestone regions
but scattered in semi-brackish ponds along the coast. An hour or two was spent
along the borders of this pond which ended up not far from the coastal dunes,
where several members of the party had their first glimpse of dune plants such
as the ever-present Euphorbia polygonifolia, sea-rocket (Cakile), seaside gol-
denrod, (Solidago sempervirens), Artenusia caudata, and the silvery-leaved
A. Stelleriana, which is commonly known as Dusty Miller.

170

ALEXANDER AND SVENSON: FIELD TRIP 171

Our transportation had been very carefully arranged by Dr. Small, and we
caught here the bus going southward to Seaside Park where we were to stop
for the night. After lunch, again through the careful planning of Dr. Small,
we went by automobile southward to Island Beach, one of the wildest places
on the New Jersey coast. This area forming the northern barrier-beach of
Barnegat Bay is many miles in length, and since it has been kept under private
ownership it is still relatively undisturbed. The dunes on the oceanside were
especially colorful with carpets of Hudsonia tomentosa, the yellow flowers pro-
jecting only an inch or so above the shifting sand. Here the prize find was
Carex macrocephala, now to be called Carex Kobomugi. The staminate and
pistillate plants are separate in the species, which forms deep-rooted mats in
the shifting dunes. Except for a station at Cape Henry, Virginia, it is not other-
wise known on the Atlantic coast; its presence is undoubtedly due to marine
shipping. Crossing to the bayward side all of our party were greatly pleased
with the large trees of various sorts which had been dwarfed and cut into fan-
tastic shapes by the wind. Here were junipers, hollies with trunks a foot or so
in diameter, splendid examples of the southern red oak (Quercus falcata)
which reaches its northern limit at about this area, and large patches of our
native cactus (Opuntia compressa).

Some of us were even more interested in the vast and variable numbers of
blueberries which filled the bushes in the damp hollows. Some of these hollows
had sphagnum with the pink orchid (Pogonia ophioglossoides) and in one of
the little depressions were plants of the smallest of the bladderworts, Utri-
cularia cletstogama. In all these hollows there were also plenty of mosquitoes.
This long tongue of land is only a few hundred yards in width and the sheltered
bayside was soon reached. Here just above the high-water mark were vast
rows of the so-called ditch grass (Ruppia maritima), cast up by the tide, and
just one fragment of the related Zanmichellia was found. Along these beaches
were numerous plants of the sow thistle (Sonchus arvensis) with attractive
large yellow flowers, a species not common in our region. The salt marsh just
to the southward was investigated by some of the members, in spite of the
mosquitoes, and here were found numerous clumps of Kosteletzkya virginica, a
mallow characteristic of salt marshes and reaching its northern limits on Long
Island and the Hackensack Meadows. By this time some of the members of the
group had become isolated in various blueberry thickets and others were al-
ready beginning the homeward journey of three or four miles to Seaside Park.
Among the interesting plants along the road were several clumps of roses of
which the identity has not yet been established. In one of the roadside ditches
were found two clumps of purple loosestrife, Lythrum Salicaria, hitherto un-
reported from this region.

172 TORREYA.

We spent the evening and night at Seaside Park (this closely built-up
town is on the seashore but none of us was able to find any trace of a “park”’).
The town is connected with the mainland by a railroad which runs over a
trestle across Barnegat Bay. It was originally planned to reach Toms River
by this railroad, but our director of transportation, Dr. Small, had found that
a motorboat could be obtained for a little more than the train fare.

At an early hour on Saturday morning, under a threatening sky, we were
embarked for Barnegat Landing, some four or five miles across the bay to the
westward, with a walk of five miles ahead for Toms River. Shortly after leav-
ing the boat it began to rain in earnest, but this rain proved to be only a shower
and the weather soon partially cleared. After we had disembarked, our road
led through salt marshes and finally to the sandy pine woods characteristic of
the pine barrens. Nothing striking was seen in these salt marshes; but at the
upper margin was a good stand of Rynchospora Torreyana, a species which is
not too abundant, and well-marked clumps of Eleocharis ambigens, the repre-
sentative of Eleocharis palustris along the southern coastal plain. As we leit
the salt marsh area the rain had stopped and we visited bogs with cotton grass
(Eriophorum), pitcher plants, and Calopogon on the way. Some of the party
stopped to browse over a burned area which was studded with Arenaria
caroliniana and Lobelia Canbyi, Liatris gramintfolia not yet in flower, and the
five coastal Eupatoriums, E. album, E. hyssopifolium, E. leucolepis, E. ver-
benaefolium and E. rotundifolium, and a long discussion was held over the
differentiation of these species. The oaks, especially possible hybrids, were the
subject of a good deal of argument, as was also the question of whether Pinus
rigida could always be determined from P. echinata by the character of the
bark. Both species of pine were here in approximately equal numbers. Probably
the most interesting discovery of the whole trip was that of Oenothera rhombi-
petala in a vacant lot at River Bank; this species is reported in Gray’s Manual
as being known from Indiana to Minnesota, Nebraska, and Texas. Our five
mile walk having been completed without much rain, we landed in the village
of Beachwood in time for lunch and a heavy downpour. Since our party now
numbered about twenty-six we pretty nearly cleaned out the eating facilities
of the village. A few who had important business in New York left the group
at this point, but the rest of us proceeded in a bus southward to the botanical
stamping ground of Forked River, and especially to the middle branch where
there is a bus-stop bearing the name of Ostrom. From here it was only a short
walk down to the river. Our principal plant of interest was the curly grass
(Schizgaea pusilla), a small fern which has always been the most interesting
single attraction of the barrens. Although one may know the exact location of
the plant from past experience, it is not always easy to find. This was true in
the present case, but the tiny plants were finally located in little hollows among

ALEXANDER AND SVENSON: FIELD TRIP 173

the Dendrium bushes, associated with Lycopodium and the orange milkwort
(Polygala lutea). In another location to the south the plants grew adjacent to
Pyxidanthera and Drosera filiformis in an open pathway where there was a
slight accumulation of sphagnum moss. The flora along the margin of the river
was as brilliant as any of us had ever seen in the pine barrens, and the slightly
cloudy weather tended to enhance the golden flowers of Lophiola and Narthe-
cium americanum, both now in full bloom. In shallow water there was an ex-
panse of yellow bladderwort (Utricularia cornuta), with little islands formed
entirely of red-leaved sundew (Drosera intermedia). Floating in the deeper
water were many colonies of Utricularia fibrosa and U. macrorrhiza. Here the
pitcher plant (Sarracenia) filled up shallow coves in unbelievable abundance,
but flowering time had long passed. It was with regret that we plodded back
a mile or so to the bus-station, since we all felt that the region could have
stood a couple of days’ exploration at the least; but our walk was somewhat
enlivened by the large number of stray species, such as Polygonum cuspidatum,
which are now appearing on the roadside rubbish-piles, characteristic of so
many of our highways.

While the bus and train took us toward New York, our party became
smaller as the members took their various ways home. Headlines in the news-
papers of fellow passengers reminded us of the sterner events in the world at
large, and made us appreciate all the more the respite we had enjoyed of a
few days in which to dwell upon the botanical achievements of the last seventy-
five years, and the opportunity to visit again some of the favorite collecting
grounds in the range of the Torrey Club. Thus, drew to its close, the Seventy-
fifth Anniversary Celebration of the Torrey Botanical Club.

Voi. 43 TORREYA DEcEMBER 1943

ACTIVITIES OF THE CLUB

May To NoveMBER 1943

May 19. Meetinc IN SCHERMERHORN EXTENSION, COLUMBIA UNIVERSITY.
The meeting was called to order by the first Vice-President, Dr. Seaver, at 3:30
p.m. Attendance: 24. The minutes of the preceding meeting were approved. The scien-
tific program was presented by Mr. Louis P. Flory of the Boyce Thompson Institute,
who spoke on “Color Photography,” discussing the problems of equipment, exposure,
and lighting in color photography. He illustrated his talk with sample slides. The meet-
ing adjourned at 4:45 p.m.
Honor M. HoLitiIncHuURST,
RECORDING SECRETARY.

June 5. Fretp Tri to the Brooklyn Botanic Garden for a study of exotic trees and a tour
of the herb garden. Leader: Miss G. Elizabeth Ashwell. Attendance: 2.
June 6. Fretp Trip along Appalachian Trail near Southfields, N. Y. The “finds” were
Betula papyrifera and Orobanche uniflora. Leader: Mr. G. G. Nearing. Attendance: 2.
June 12. Frevp Trip to Egbertville, Staten Island. Leader: Mr. Charles Ericson. Attend-
ance: 3.
June 20. Frevp trip to Montclair Heights, N. J. Joint outing with the Newark Museum
Nature Club. Leader: Prof. Oliver P. Medsger. Attendance: 39.
June 26. Fietp Trip to the New York Botanical Garden to see the laboratory and field
work of the leader, Dr. A. B. Stout, Director of Laboratories. Attendance: 3.
Juty 4. Frecp Trip to Arden, N. Y. for fungi. Most prized catch was Hygrophorus psitia-
cinus, one of the few fungi with a green color. Leader: Mr. F. R. Lewis. Attendance 7.
Jury 11. Frey Tr along Stony Brook Trail, Sloatsburg, N. Y. for lichens and fungi,
both of which were fairly abundant. Leader: Mr. G. G. Nearing. Attendance: 10.
Juty 17. Frey Trp to the home of Mr. W. H. Dole, our leader, to see many species of
native and introduced ferns as garden plants. Attendance: 33.
Jury 25. Frevp Trip along the Kakiat Trail at Tuxedo, N. Y., for lichens, fungi, and gen-
eral botany. Leader: Mr. G. G. Nearing. Attendance: 18.
Avucust 1. Fietp Trip to Sloatsburg, N. Y. A successful search for Boleti. Leader: Mr.
F. R. Lewis. Attendance: 9.
‘Aucust 8. Fievp Tri to climb Schunemunk Mt, Washingtonville, N. Y. Leader: Dr.
Alexander V. Tolstoouhov. Attendance: 5.
Aucust 14. Frecp Tri to the vicinity of Midvale, N. J., for fungi. Leader: Mr. F. R.
Lewis. Attendance: 2.
Aucust 15. Fretp Tri to Glen Cove, L. I., for fossils and general botany. Leader: Mr.
James Murphy. Attendance: 9.
Aucust 22. Fretp Tre to Mt. Vernon and the Bronx, for general flora, asters and golden-
rods in particular. Due to confusion about the assembly point, two trips were held. Mrs.
Mary Holtzoff, the scheduled leader, had 6 present, and Mr. Joseph Monachino led a
group of 11. Each group reported a satisiactory outing.

Aucust 29. Fretp Tre to Harmon, N. Y., for fungi. The dry season reduced the number
found materially. Leaders: Mr. F. R. Lewis and Mr. A. D. Mebane. Attendance: 3.
SEPTEMBER 11. Fretp Trr to Butler, N. J., for fungi, but the weather had continued dry
and fungi were less than scarce. However, Russula elegans was found. Leader: Mr. F. R.

Lewis. Attendance: 3.

. 174

ACTIV IDI ES Oh Vir CLUB 175

SEPTEMBER 12. FIELD Trip to Preakness Hills, N. J., for lichens, fungi, and general botany.
Stereocaulon pileatum was found. This is believed to be the first record for New Jersey
of this species which is usually collected in the Adirondack or White Mountains.
Leader: Mr. G. G. Nearing. Attendance: 17.

SEPTEMBER 18. FIELD Trip to Richmond Valley, Staten Island, N. Y. Several hybrid oaks
were seen and the general botany observed. Leader: Mr. W. T. Davis. Attendance: 25.

SEPTEMBER 29. FIELD Trip to the Boyce Thompson Arboretum led by Mr. J. H. Beale,
Superintendent. Attendance: 10.

SEPTEMBER 26. Fietp Trip to Mineola, L. I., N. Y., for Myxomycetes. Still too dry. Leader :
Mr. Robert Hagelstein, Honorary Curator of Myxomycetes at The New York Botan-
ical Garden. Attendance: 6.

SEPTEMBER 26. FIELD Trip to Van Cortlandt Park, Bronx, N. Y. Fraxinus nigra and F.
pennsylvanica were found. They are not often seen in this vicinity. Leader: Dr. A. H.
Graves of Brooklyn Botanic Garden. Attendance: 18.

Octoser 2. FieELD Trip to Grassy Sprain region, Yonkers, N. Y., for fungi. The leader,
Dr. M. Levine, reported a “perfect trip.” Attendance: 3.

Ocroser 3. Fietp Trip to Point Pleasant vicinity, N. J. Species attracting most attention
were Gentiana saponaria, Polygala Nuttallii and P. cruciata, Jasione montana, Bar-
tomia virginica, and a species of Sabatia. Leader: Mr. V. L. Frazee. Attendance: 3.

OcroBer 5. MEETING AT THE BROOKLYN BOTANIC GARDEN.

The meeting was called to order by the President, Dr. Robbins, at 8:15 p.m. Attend-
ance: 34. The minutes of the meeting of May 19th were approved. Twenty-two persons
were elected to annual membership and seven to associate membership. It was voted to
invest another $10,000 of the capital of the Club in war bonds. The collecting experiences
of the Club members provided the scientific program of the evening. These experiences
ranged from collecting on field trips or by proxy to working in victory gardens; from
identifying an uncommon plant to research in the field of rubber. By the conclusion,
a picture of the varied fields of interest of the Torrey Club members had been pre-
sented. The meeting adjourned at 9:40 p.m. and refreshments were served by members
of the Garden staff.

Honor M. Ho.tiincHurst,
RECORDING SECRETARY.

Ocroser 10. Frerp Trip to Richmond, S. I., N. Y., for general flora of brookside, old
fields, and salt marsh. Leader: Miss Hester M. Rusk of the Brooklyn Botanic Garden.
Attendance: 15.

Ocroser 17. FrELp Trip to the Brooklyn Botanic Garden for the study of coniferous plants.
Leaders: Drs. A. H. Graves and Alfred Gundersen of the Garden staff. Attendance: 11.

Octoser 20, 1943. MretInc at THE NEw York BotanicAL GARDEN.

The meeting was called to order by the President, Dr. Robbins, at 3:30 p.m. Attend-
ance: 37. The minutes of the preceding meeting were approved. A Memorial Tribute
to the late Dr. C. Stuart Gager was read by Dr. Dodge, chairman of the Memorial
Committee :

October 20, 1943.
It is with profound sorrow and a realization of a great loss to our organization
that the Torrey Botanical Club records here the death of Doctor Charles Stuart

Gager, who died August 9, 1943.

Doctor Gager was elected to membership in the Club October 25, 1905. He had
served the Club with high honor and distinction, not only on committees which had
to do with formulating plans and policies, but also as Recording Secretary for three

176

UD (QUIS TR TE NEN

years and delegate of the Club for several years on the Council of the New York
Academy of Sciences and to the Council of the American Association for the Ad-
vancement of Science. He was Vice-President for fourteen years, from 1917 to
1941, and served the Club well as its President during the year just previous to his
death.

In recent years it had been the custom of the Club to hold its first fall meeting
at the Brooklyn Botanic Garden, at which time members were given an opportunity
to report on their work during the summer period. Those who attended these meet-
ings will long remember the cordiality and sincerity of his greeting which always
left one with the impression that in him one had a very warm personal friend.

As a token of appreciation of the importance of his contributions to our knowl-
edge of plants and recognition of his administrative abilities, and also for his ex-
ample of right living, it is directed that this memorial be published with the minutes
of this meeting and a copy sent to members of the bereaved family.

(Signed) Sam F. TRELEASE
P. W. ZIMMERMAN
ARTHUR H. GRAVES
B. O. Donce, Chairman.

After Dr. Dodge so moved, the Memorial was accepted by a rising vote of the Club
members.

The scientific program was presented by Dr. Michael Levine, who spoke on “The

combined effects of colchicine and x-rays on onion root tips.”

Fifteen series of experiments were made in which six to forty onions (Alliwn
cepa var. Yellow Globe or var. Brigham Yellow Globe) were used in each. The
bulbs were selected for their uniformity of weight, size and freedom from fungus
diseases. The bulbs were placed in water for periods of three to twelve days to in-
sure an adequate number of roots. The bulbs were then placed in a 0.01 per cent
aqueous solution of colchicine and after 6,18,24,36,48,72,96,125, or 140 hours of
exposure were removed and washed in running water. After each given exposure, be-
ginning with the 18 hour treatment the bulbs were divided into two groups of equal
number. The first group was returned to fresh water, the second group was ex-
posed to x-rays. A third group, not treated with colchicine, was irradiated simul-
taneously with the second group; both groups were then returned to water. A fourth
group of bulbs was kept in water. The x-ray treatment consisted of a single exposure
for 11 to 30 minutes during which time 900, 1500 or 3000 roentgen units (r) were
delivered.

The roots of the four groups of bulbs were examined daily and photographed at
frequent intervals. Selected root-tips from all the bulbs were prepared for micro-
scopical examination. Root-tips exposed to colchicine for 72,96,125, or 140 hours and
irradiated with 900 r when returned to water failed to grow. Root-tips exposed to
colchicine for 24 to 48 hours and irradiated with 900 r showed temporary growth
inhibition and resumed growth as indicated by the prolongation of the tips below the
swellings induced by the colchicine. With larger doses of x-rays, 1500 r and 3000 r,
roots colchicinized for 36 to 48 hours failed to resume growth for 14 to 21 days after
their return to water.

The microscopical examination of these arrested tissues showed progressive
coagulation and destruction of the nuclear materials of the cells in the root-tips.
The root-tips colchicinized only were studied concurrently but showed complete re-
covery when returned to water. The roots irradiated, only showed temporary arrests
of growth. With the higher doses of x-rays some injury was noted but growth was
halted temporarily.

Acenaphthene used in lieu of colchicine had no effect on the activity of the x-rays.
Roots so treated behaved like those non-chemically treated.

The combined effect of colchicine and x-rays was also studied on the growth of
leaves of the onion of the Brigham Yellow Globe variety. The leaves of the col-
chicinized bulbs showed little growth after irradiation with 3000 or 1500 r. While
the plants x-rayed only showed some leaf growth but less than that which occurred in
normal or colchicinized bulbs. The latter two groups showed little difference between
them.

ACTIVITIES OF THE CLUB 177

These observations led to the conclusion that colchicine sensitizes the formative
embryonic tissue of the root to x-rays. The influence was not determined solely
by the division phase of the nucleus. The resting nuclei as well as the dividing ones
seemed to be affected. The effect of the colchicine and x-rays on the dividing
nuclei was more obvious for the chromosomes in metaphase stage were clumped or
coagulated while no visible change appeared in those of the resting phase.

It appears that colchicine combined with x-rays has a definite role in cancer ther-
apy. Some tumors of known cytogenetic homogeneity should be the basis for fur-
ther study.

The meeting adjourned at 5:00 p.m. Tea was then served by members at the Garden.
Honor M. Horiincuurst,

RECORDING SECRETARY.

Octoser 24. Firtp Trip to Alpine and the Palisades, N. J. General leader: Mr. G. G.
Nearing. Assistants: for fungi, Mr. F. R. Lewis; for lichens, Mr. W. L. Dix; for
bryophytes, Dr. Holberg; for higher plants, Mr. L. E. Hand. Attendance: 20.

Octoser 31. Members were invited to participate in the annual pilgrimage of many New
York hiking clubs to Long Mountain in Palisades Interstate Park, in memory of the
late Mr. Raymond H. Torrey, who was President of the Torrey Botanical Club when
he died in 1938. ;

Dr. Small, the chairman of the Field Committee, reports that during 1943 a total
of 43 field trips were arranged. This is about one-half of the number of trips offered
in recent years. The total attendance was 485, or about one-third of that of recent
years.

NoveMBER 17. MEETING aT THE NEw York BOTANICAL GARDEN.

The meeting was called to order at 3:30 p.m. by the President, Dr. Robbins. Attend-
ance: 36. The minutes of the preceding meeting were approved. It was voted that the
Club act as host to any sectional meeting of the Botanical Society of America which
might be held in the New York area.

Dr. Matzke read the following letter from Mrs. C. Stuart Gager :

29 Linden Boulevard
Brooklyn.

Dr. Epwin B. MatzKE

Corresponding Secretary

Torrey Botanical Club

My dear Dr. Matzke,

I am deeply grateful to the Torrey Botanical Club for the high tribute paid to my
beloved husband in the Memorial recorded in the Minutes of the meeting of the Club on
October 20.

This expression of their esteem and sense of loss in his passing is most sincerely
appreciated.

Faithfully yours,
(Signed) Berroa B. GaAcEeR
NOVEMBER 14.

The first part of the scientific program was presented by Dr. Bassett Maguire, and
entitled a “Report on the 1943 Field Summer in the Great Basin.”

A general and brief description of the physiographic and vegetative character-
istics of the Intermountain Region was given. In somewhat more detail the struc-
tural and floral characters of the Deep Creek and Raft River Ranges, Utah, and
the Ruby, North Humboldt, and Santa Rosa Ranges, Nevada, were discussed.
The net results of the summer’s activities were listed as approximately 1000 numbers
and 6000 sheets collected.

178 NO RAE WEA

Mr. Robert Hulbary, the second speaker, discussed “Three Dimensional Cell Shapes
in the Differentiating Cortex of Elodea Stems.”

There were three purposes for making this study; one to determine the three
dimensional shapes of the mature cortical cells in the presence of large air spaces,
another to investigate the shapes of the cells in the apical meristem, and a third to
study changes in shape as the cortical cells differentiate.

In the cortex of Elodea (Anacharis densa Victorin) the three-dimensional shapes
of the cells in the stem cortex are influenced by the presence of large internodal
lacunae. The cortical cells are elongated parallel to the long axis of the stem,
and they contain chloroplasts and starch grains. One hundred cells from each of 27
consecutive internodes were studied to determine the number of faces per cell. Then
600 additional cells—100 from each fifth internode—were studied more intensively
for number and kinds of faces and the combinations of faces. The average number
of faces per cell for the 3300 cells was 8.79. More than one third of all the cells
(1443) were 8-hedra. Quadrilateral faces occurred more frequently than all of the
other kinds added together. In 600 cells studied more intensively only 31 different
combinations of faces were encountered. This apparent uniformity in cell shape in
Elodea stem cortex is further attested to by the fact that three of these 31 patterns
were outstandingly characteristic for the issue.

Using the method of Duchartre, the average number of faces per cell in the apical
meristem was found to be 13.88.

The large internodal air canals originate schizogenously, and they are completely
delimited at the base of the apical meristematic region. The reduction in number of
faces per cell from the apical meristem to the mature cortex and the other differenti-
ations in cell shape concomitant with the development of the internodal lacunae are
due to the cell enlargement and to cell divisions which are limited to two distinct
planes.

Following discussion of the two talks, the meeting was adjourned at 4:55 p.m. Tea and
refreshments were then served by friends at the Garden.
Honor M. HoLiincHurst,
RECORDING SECRETARY.

ADDITIONS TO THE List oF BOTANISTS IN THE FRONTISPIECE

No. 24. For C. F. Mook read P. V. Mook
No. 47. Mrs. George S. Powell

No. 48. George S. Powell

No. 49. Mrs. Robert Hagelstein

DATES OF PUBLICATION OF TORREYA, VOLUME 43

Number 1. July August 27, 1943
Number 2. December February 10, 1944

+ peg

Vo. 43

A OP RORS EEN

DECEMBER 1943

INDEX TO TORREYA—VOLUME 43

Acarospora murorum, 82

Acer rubrum, 83

Activities of the Club, 78, 174

Ailanthus, 83

Aletris farinosa, 170

ALEXANDER, E. J. & Svenson, H. K.: The
field trip to the New Jersey coast and
pine barrens, June 26-27, 1942, 170

Alisma Plantago-aquatica 46, subcordata,
45

ALLEN, C. E. 2: The evolution and deter-
mination of sexual characters in the
Angiosperm sporophyte, 6

Allium cepa, 176

Anacharis densa, 178

Angiosperm sporophyte, Evolution and de-
termination of sexual characters in, 6

Antirrhinum, 10

Arenaria caroliniana, 172

Artemisia caudata, 170; Stelleriana, 170

Asclepias lanceolata 170; rubra, 170

Asimina triloba, 61

Aspergillus mger, 124

Aster yellows, 94, 95

Astilbe japonica, 61

Bartonia virginica, 175

Benzoic acids, 102, 107, 109-113, 115

Betula papyrifera, 174

Bicknell, Eugene P., 40

Biotin, 119-121

BLAKESLEE, A. F., 3, 169

Botanists attending the 75th Anniversary
Celebration of the Torrey Club 1, 178

Britton, Nathaniel L., 42, 43

Brown, Addison, 41

Bryonia dioica, 8

Cain, S. A. 3; Criteria for the indication
of center of origin in plant. geographical
studies, 132

Calendula, 91, 92

Campanula, calycanthema forms of, 10

Canna, 9

Carex Kobomugi, 171; macrocephala, 171

Carica Papaya, 61

Caulophyllum thalictroides, 45

Ceanothus, 135

Cell division as a problem of pattern in plant
development, 29

Cell shapes, Three dimensional, in the dif-
ferentiating cortex of Elodea stems, lec-
ture, 178 ;

Cellular behavior during regeneration, lec-
ture, 82

Chaulmoogra odorata, 58

Cheiranthus, 9

Cinchona, 59

Colchicine and x-rays, The combined effects
of, on onion root tips, lecture, 176

Cornus, The genus in North America, lec-
ture, 78; amomum, 79; canadensis, 79;
florida, 79; Kousa, 79; mas, 79, 83; ses-
silis, 79; stolonifera, 79; suecica, 79

Crepis, 137

Criteria for the indication of center of ori-
gin in plant geographical studies, 132

Crocker, WILLIAM, 3

Cronartium ribicola, 157, 158

Cucumis, sativus, 114

Cunninghamia lanceolata, 24

Cyphelium tigillare, 82

Cypripedium Calceolus 45; pubescens, 45

Datura stramonium, 114

Dawson, Ray F.: Some aspects of parasit-
ism in the mycorrhizae of shortleaf pine,
lecture, .81

Dentaria, 84

Digitalis purpurea, 60

Diphylleia cymosa, 45

Drepanocladus, Variability and distribution
of, in North America, lecture, 80

Drosera filiformis, 173; intermedia, 173

Drosophila, 14, 69, 127

Economic aspects of taxonomy, 50

Elatine americana, 170

Eleocharis ambigens, 172; microcarpa, 46;
palustris, 172

Elodea, 178

Endothia parasitica, 157

Ephedra, 19; sinica, 57, 58

Epilobium, 13

Equisetum, 23, 24, 29, 33; hyemale, 30, 31

Erica, 136

180 TORREYA

Eriocaulon septangulare, 170

Erythronium, 84

Eupatorium, oriented pith of, 5; album, 172;
hyssopifolium, 172; leucolepis, 172; ro-
tundifolium, 172; verbenaefolium, 172

Euphorbia polygonifolia, 170

Evolution and determination of sexual char-
acters in the Angiosperm sporophyte, 6

False blossom disease, 87-95

Ficus, 57

Field trips in 1943, Leaders:
AppotT, Mrs. RicHarp M., 84
ASHWELL, G. ELizapetu, 174
Beate, J. H., 175
Connon, NELLIE L., 82, 84
Davis, W. T., 175
Dixs, Weel) 177
Dott, W. H., 174
Ericson, CHARLES, 174
Frazer, V. 1, 175
Graves, A. H., 84, 175
GUNDERSEN, ALFRED, 84, 175
HAGELSTEIN, Ropert, 175
Hanp, L. E., 85, 177
Horserc, Dr., 177
Hottzorr, Mrs. Mary, 85, 174
Husk, W. M., 85
JoHNSON, JuLtus, 85
Lewis, F. R., 85, 174, 177
Mesange, A. D., 174
Menscer, O. P., 85, 174
MoNACHINO, JOSEPH, 174
MurpHy, JAMes, 174
NEARING, G. G., 82, 84, 85, 174, 175, 177
Rusk, Hester M., 175
SMatt J. A., 84
TotstoouHoy, A. V., 174
Witey, Faripa A., 84

Field trips in 1943, Locations:
Arden, N. Y., 174
Boyce Thompson Arboretum, 175
Branchville, N. J., 85
Brooklyn Botanic Garden, 84, 174, 175
Butler, N. J., 174
Central Park, New York, 82
Glen Cove, L. I., 174
Grassy Sprain, Yonkers, 175
Harmon, N. Y., 174
Haskell, N. J., 85
McLean Woods, New York, 85

Midvale, N. J., 174
Mineola, L. I., 175
Montclair Heights, N. J., 174
Mt. Vernon and The Bronx, 174
New York Botanical Garden, 84, 174
New York Zoological Park, 82
Palisades Interstate Park, 84, 177
Point Pleasant, N. J., 85, 170, 175
Preakness Hills, N. J., 175
Raritan River, 84
Ridgewood, N. J., 85
Schunemunk Mountain, 174
Silver Lake, White Plains, 84
Sloatsburg, N. Y., 174
Southfields, N. Y., 174
Springdale, N. J., 82
Staten Island, 174, 175
Surprise Lake, Summit, N. J., 84
Tuxedo, N. Y., 174
Van Cortlandt Park, N. Y., 175
Fir, Douglas, 83
Fiory, L. P.: Color photography, lecture,
174
Formative influences and comparative ef-
fectiveness of various plant hormone-like
compounds, 98
Fragaria, 11, 14
Franseria dumosa, 139
Fraxinus nigra, 175; pennsylvanica, 175
Freycinetia, 56
Fungi, The importance of taxonomic stud-
ies of, 65

Gacer, Bertua B., 177

Gacer, C. Stuart, 3, 175

Gaylussacia, 135

Genetics, the unifying science in biology,
126

Gentiana saponaria, 175

Geography, relation to modern taxonomy,
44

Geranium, 10, 13

Geum, 10

Ginkgo, 19; biloba, 24

Gieason, H. A. 3, 84; Contributions of the
Torrey Botanical Club to the develop-
ment of taxonomy, 35

Gleichnia, 23

Gratiola aurea, 170

Gray, Asa, 35, 36, 44, 56

INDEX 181

Gymnosporangium juniperi-virgiuuanae,
158; sabinae, 158
Gynocardia odorata, 58

Haphazard as a factor in the production of
tetrakaidecahedra, 4

Hazen, Tracy E., 82, 83

Hevea, 59

Hieracium excellens, 13

Hippuris vulgaris, 19, 24

Hollick, Arthur, 41

Hormones, Animal, affecting growth and
several effects of single hormones, 96

Hormone-like compounds, Formative in-
fluences and comparative effectiveness of,
98

Hutpary, Ropert: Three dimensional cell
shapes in the differentiating cortex of
Elodea stems, lecture, 178

Hydnocarpus Alcalae, 58, 59; castaneus, 58;
Hutchinsonu, 58; Kursti, 58; subfalcata,
58; Woodii, 58, 59

Hydrocotyle uwmbellata, 170

Hygrophorus psittacinus, 174

Aymenophyllum, 23

Indians, Local plants used by, Lecture, 79
Inferiority complexes in plants, lecture, 82

Jasione montana, 175
Juncus, stellate 12-rayed cells of, 5

Karine, J. S., 2

Kern, F. D. 3; The importance of taxo-
nomic studies of the fungi, 65

Kosteletzkya virginica, 171

KunkeL, L. O. 3; Viruses in relation to
the growth in plants, 87

Larrea tridentata, 139

Leaf-stem relationships
plants, 16

Lesquerella, 135, 140, 148

LevINE, MicHaet, 175; The combined ef-
fects of colchicine and x-rays on onion
root tips, lecture, 176

Lewis, F. T. 2; Haphazard as a factor in
the production of tetrakaidecahedra, 4

Liatris graminifolia, 172

Linum, 13, 25

Lobelia Canbyi, 172

Lonicera fragrantissima, 83

Lophiola, 173

Lychnis, 14; alba, 8; dioica, 8

in the vascular

Lycopodium, 173; annotinum, 22; cernum,
22; clavatum, 22; complanatum, 22; var.
flabelliforme, 22; inundatum, 22; lucid-
ulum, 21; obscurum, 22-25; sabinaefoliwm,
22; Selago, 21

Lygodium, 23

Lythruwm Salicaria, 171

Macuirre, Bassetr: Report on the 1943
field summer in Great Basin, lecture, 177

Matzke, E. B., 2, 82, 83

Medinilla, 56

Meetings of the Club in 1943:

Jan. 5, 78 Apr. 21, 84
Jan. 20, 78 May 11, (84
Feb. 2, 79 May 19, 174
Feb. 17, 79 Oe 5, 75
Mar. 2, 81 Oey Abs 7
Mar. 17, 82 Nov. 17, 177
Apr. 6, 83

Mercurialis, 11

Merritz, E. D., 3; Some economic aspects
of taxonomy, 50

Mertensia, 84

Mucor ramannianus, 124

Mycorrhizae of shortleaf pine, lecture, 81

Myriophyllum humile, 170; tenellum, 170

Naphthalene compounds, 99, 100

Naphthoxy compounds, 99-101, 114, 115

Narthecium americanum, 173

Naytor, Ernest: Problems of cellular be-
havior during regeneration, lecture, 82

Nephrolepis, 23-25

Nicandra physalodes, 101, 106, 107

Nicotiana, 13; glutinosa, 91, 93

Oenothera, 9, 127; longipedicellata, 170;
rhombipetala, 172

Officers of the Club for 1943, 78

Ononts, 135

Opuntia compressa, 171

Orobanche uniflora, 174

Oryza, 10

Pandanus, 56

Panicum meridionale, 46

Papaver, 9

Peach yellows, 94, 95, 167

Phenoxy compounds, 101-108, 114, 115
Phleum pratense, 10

Photosynthesis in bacteria, lecture, 80

182 ay ORRGRUECOviE

Phycomyces, 117, 123, 124

Physalis alkekengi, 87

Physcia venusta, 82

Phytopathology, 1867-1942, 155

Phytophthora cinnamomi, 124; infestans,
156

Pine, parasitism in the mycorrhizae of,
lecture, 81; Austrian, 83; shortleaf, 81;
white, 83

Pinus echinata, 172; rigida, 172; Strobus,
24

Plant development, Cell division as a prob-
lem of pattern in, 29

Pogonia ophioglossoides, 171

Polygala cruciata,-175; lutea, 170, 173;
Nuttalli, 175

Polygonum cuspidatum, 173

Potamogeton pectinatus, 170

Psaronius, 23

Pscudotsuga, 24, 25

Pteridium aquilinum, 45; latiusculum, 45

Puccinia coronata, 158, 162; glumarum, 165;
graminis, 68, 155, 157, 159, 160, 162; ru-
bigo-vera tritici, 160; subnitens, 160

Pyridoxine, 118

Pythium butleri, 123

Quercus Cerris, 83; falcata, 171; phellos,
170

Ranunculus, 47; sceleratus, 170

Recorp, SAMUEL: How woods are identi-
fied, lecture, 84

Reep, G. M., 3; Phytopathology, 1867-1942,
155

Rhododendron, calycanthema forms of, 10

Ricxett, H. W.: The genus Cornus in
North America, lecture, 78

Rrppie, Oscar, 3; Animal hormones affect-
ing growth and the several effects of
single hormones, 96

Roggtns, W. J., 3; Plants need vitamins
too, 116

Rubus, 12

Rumex, 14

Ruppia maritima, 171

Rusby, Henry H., 42, 43

Russula elegans, 174

Rynchospora macrostachya, 46; Torreyana,

WZ
Si abatia, 175

Salix, 11

Sararanga, 56

Scalesia, 48

Schizaea pusilla, 172

Scleria reticularis, 46

Sclerotium rolfsi, 124

Seleginella, 22, 24, 25

Senecia, 136

Sequoia, 24

SHULL, G. H., 3; Genetics, the Unifying
science in biology, 126

Silene, 8, 9, 11

Sinnott, E. W., 2; Cell division as a prob-
lem of pattern in plant development, 29

Smilacina bifolia, 45; var. canadensis, 45;
var. kamtschatica, 45

Sonchus arvensis, 171

Spinacia, 11

Spiraea, 61

Stereocaulon pileatum, 175

Stout, A. B., 174; Dichogamy in relation to
reproduction, lecture, 84

Streptocarpus, 13

SveENsoN, H. K., 3; Modern taxonomy and
its relation to geography, 44; Plants of a
Long Island pond, lecture, 84

SvENSON, H. K. & ALEXANDER, E. J.: The
field trip to the New Jersey coast and
pine barrens, 170

Swirt, F. R.: Treating yeast plants as in-
dividuals, lecture, 79

Syringa vulgaris, 17, 19

Taraktogenos, 59; Kurzi, 58

Taxonomic studies of the fungi, Importance
of, 65

Taxonomy and its relation to geography,
44

Taxonomy, development of, Contributions
of Torrey Botanical Club to, 35

Taxonomy, some economic aspects of, 50

Taxus baccata, 17 ;

Tetrakaidecahedra, Haphazard as a factor
in the production of, 4

Thiamine, 117, 118, 123, 124

Tilletia, 161; levis, 160; tritici, 160

Tomato, 107, 109, 113

Torrey Botanical Club, Contributions of, to
the development of taxonomy 35; Offi-
cers for, 1943, 78

Torrey, John, 2, 35, 36, 39, 44, 52-54

MINE IB) 18, 2x 183

Tradescantia, 139; canaliculata, 141; hir-
sutiflora, 142; occidentalis, 141; ozarkana
142

Trichocereus spachianus, 21, 24

Ulmus americana, 83; campestris, 83

Uredo, 66

Uromyces, 66

Urtica cannabina, 11

Ustilago hordei, 77, 160; tritici, 160; vio-
lacea, 160; zeae, 160

Utricularia cleistogama, 171; cornuta, 173;
fibrosa, 173; gibba, 170; macrorrhiza, 173

Vaccinium arkansanum, 134; australe, 134;
corynibosum, 134; simulatum, 134

Vascular plants, Leaf-stem relationships in
16

Vinca rosea, 92

Viola Brittoniana, 170

Viruses in relation to the growth of plants,
87

Vitamins, Plants need for, 116

Vitis, 11

Wetmore, R. H., 2; Leaf-stem relationships
in the vascular plants, 16

Wauatey, W. G.: Inferiority complexes in
plants, lecture, 82

Winoxkur, Morris: Photosynthesis in bac-
teria, lecture, 80

Wirtrock, G. L.: Local plants used by the
American Indians, lecture, 79

Woods, How identified, lecture, 84

WYNNE, FRANcEs E.: Variability and dis-
tribution of Drepanocladus in North
America, lecture, 80

X-rays and colchicine, The combined ef-
fects of, on onion root tips, lecture, 176

Yeast plants, treated as individuals, lecture,

Zea mays, 69

ZIMMERMAN, P. W. 3; The formative in-
fluences and comparative effectiveness of
various plant hormone-like compounds,
98

THE TORREY BOTANICAL CLUB

Council for 1943

Ex officio Members

William J. Robbins Edwin B. Matzke Michael Levine
John S. Karling Honor M. Hollinghurst John A. Small
Fred J. Seaver W. Gordon Whaley Bernard O. Dodge
Lela V. Barton Harold W. Rickett

Elected Members

1941-1943 : 1942-1944 ~ 1943-1945
John H. Barnhart J. M. Arthur — . Charles A. Berger
R. C. Benedict W. J. Bonisteel Clyde Chandler
Helen M. Trelease Arthur H. Graves Albert E. Hitchcock
P. W. Zimmerman Sam F. Trelease Roger P. Wodehouse

Committees for 1943
ENDOWMENT CoMMITTEE

Clarence Lewis, Chairman Henry de la Montagne
J. Ashton Allis Helen M. Trelease
Caroline C. Haynes

ProcraMm COMMITTEE

Edwin B. Matke, Chairman (ex officio) A. B. Stout
Charles A. Berger W. Gordon Whaley
Arthur H. Graves P. W. Zimmerman

Honor M. Hollinghurst

FIELD COMMITTEE

Joun A. Smarty, Chairman

Edward J. Alexander Inez M. Haring Rutherford Platt
Vernon L. Frazee Michael Levine Daniel Smiley, Jr.
Eleanor Friend H. N. Moldenke Henry K. Svenson
Alfred Gundersen james Murphy Farida A. Wiley

- Robert Hagelstein G. G. Nearing Ellys B. Wodehouse

{
Locat Frora COMMITTEE

Epwin B. Matzke, Chairman

Edward J. Alexander Robert L. Hulbary Hester M. Rusk

H. Allan Gleason James Murphy Ora B. Smith
Arthur H. Graves William J. Robbins P. W. Zimmerman
Crypitogams

Ferns and Fern Allies: R. C. Benedict, W. Herbert Dole, N. E. Pfeiffer
Mosses: E. B. Bartram

‘Liverworts: A. W. Evans, E. B. Matzke

Freshwater Algae: H. C. Bold

Marine Algae: J. J. Copeland

Fungi: A. H. Graves, J. S. Karling

Lichens: J. W. Thomson, Jr.

Myzomycetes: R. Hagelstein

PUBLICATIONS EXCHANGE COMMITTEE

Edwin B. Matzke, Chairman (ex officio) Amy L. Hepburn Lazella Schwarten

OTHER PUBLICATIONS

OF THE

TORREY BOTANICAL CLUB>

(1) BULLETIN

A journal devoted to general botany, established in 1870 and pub-
lished bi-monthly at present. Vol. 69, published in 1942, contained 711
pages of text and 44 full page plates. Price $6.00 per annum. For
Europe, $6.25.

In addition to papers giving the results of research, each issue con-
tains the INDEX To AMERICAN BoTANICAL LITERATURE—a very compre-
hensive bibliography of current publications in American botany. Many
workers find this an extremely valuable feature of the BULLETIN.

Of former volumes, 24-69 can be supplied separately at $6.00 each;
certain numbers of other volumes are available, but the entire stock of
some numbers has been reserved for the completion of sets. Single copies
(75 cents) will be furnished only when not breaking complete volumes.

(2) MEMOIRS

The Memorrs, established 1889, are published at irregular intervals.
Volumes 1-18 are now completed. Volume 17, containing Proceedings
of the Semi-Centennial Anniversary of the Club, 490 pages, was issued
in 1918, price $5.00.

Volume 18, no. 1, 108 pages, 1931, price $2.00. Volume 18, no. 2,
220 pages, 1932, price $4.00. Volume 18 complete, price $5.00.

Volume 19, no. 1, 92 pages, 1937, price $1.50. Volume 19, no. 2, 178
pages, 1938, price $2.00.

(3) INDEX TO AMERICAN BOTANICAL LITERATURE

Reprinted monthly on cards, and furnished to subscribers at three
cents a card. .
Correspondence relating to the above publications should be ad-
dressed to
W. Gorpon WHALEY,
Barnard College,
Columbia University,
New York, N. Y.°

WOO Ut

5 00310 6299

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